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SMITHSONIAN  INSTITUTION 
UNITED  STATES  NATIONAL  MUSEUM 


PROCEEDINGS 


UNITED  STATES  NATIONAL  MUSEUM 


VOLUME    103 

NUMBERS  3311-3337 


UNITED  STATES 

GOVERNMENT  PRINTING  OFFICE 

WASHINGTON  :  19S6 


ADVERTISEMENT 

The  scientific  publications  of  the  National  Museum  include  two 
series,  known,  respectively,  as  Proceedings  and  Bulletin. 

The  Proceedings,  begun  in  1878,  are  intended  primarily  as  a  medium 
for  the  publication  of  original  papers,  based  on  the  collections  of  the 
National  Museum,  that  set  forth  newly  acquired  facts  in  biology, 
anthropology,  and  geology,  with  descriptions  of  new  forms  and 
revisions  of  limited  groups.  Copies  of  each  paper,  in  pamphlet 
form,  are  distributed  as  published  to  libraries  and  scientific  organiza- 
tions and  to  specialists  and  others  interested  in  the  different  subjects. 

The  dates  at  which  these  separate  papers  are  published  are  recorded 
in  the  tables  of  contents  of  each  of  the  volumes. 

The  present  volume  is  the  hundred  and  third  of  this  series. 

The  Bulletin,  the  first  of  which  was  issued  in  1875,  consists  of  a 
series  of  separate  publications  comprising  monographs  of  large 
zoological  groups  and  other  general  systematic  treatises  (occasionally 
in  several  volumes),  faunal  works,  reports  of  expeditions,  catalogs  of 
type  specimens,  special  collections,  and  other  material  of  similar 
nature.  The  majority  of  the  volumes  are  octavo  in  size,  but  a  quarto 
size  has  been  adopted  in  a  few  instances  in  which  large  plates  were 
regarded  as  indispensable.  In  the  Bulletin  series  appear  volumes 
under  the  heading  Contributions  from  the  United  States  National 
Herbarium,  in  octavo  form,  published  by  the  National  Museum  since 
1902,  which  contain  papers  relating  to  the  botanical  collections  of  the 
Museum. 

Remington  Kellogg, 
Director,  United  States  National  Museum. 


CONTENTS 

Pages 

Abbott,  K.  Tucker.  Review  of  the  Atlantic  periwinkles 
Nodilittorina,  Echininus,  and  Tedarius.     Figures    55-57. 

Published   March   23,    1954 449-464 

Arnett,  Ross  H.,  Jr.  Beetles  of  the  oedemerid  genus 
Vasaces    Champion.     Figure    13.     Published    April    30, 

1953 87-94 

New  species:  Vasaces  linearis,  V.  knulli,  V.  maculatus,  V.  elon- 
gaius, 

.     A    review    of    the    beetle    family   Cephaloidae. 

Figure  20  and  plate  5.     Published  May  15,  1953   ....    155-161 

.     Supplement   and   corrections   to   J.    A.   Hyslop's 

Genotypes  of  the  elaterid  beetles  of  the  world.     Published 

April  14,  1955 599-619 

Banner,  Albert  H.  A  supplement  to  W.  M.  Tattersall's 
Review  of  the  Mysidacea  of  the  United  States  National 

Museum.     Published  July  8,  1954 575-583 

Berger,  Andrew  J.  On  the  anatomy  and  relationships 
of  glossy  cuckoos  of  the  genera  Chrysococcyx,  Lampro- 
morpha,  and  Chalcites.    Figures  69-71.    Published  January 

19,  1955 585-597 

Blake,  Doris  Holmes.  The  chrysomelid  beetles  of  the 
genus  Strabala  chevrolat.     Figure  17.     Published  June  5, 

1953 121-134 

New  species:  Strabala  rotunda,   S.  acuminata,  S.  colombiana,  S. 

trinitatis. 
New   subspecies:    Strabala    ambulans    jamaicensis,  S.  a.  puerto- 
ricensis,  S.  rufa  floridana,  S.  acuminata  teapensis,  S.  a.  costa- 
ricensis. 

Carriker,  M.  a.,  Jr.  Studies  in  Neotropical  Mallophaga, 
XI:  Bird  lice  of  the  suborder  Amblycera,  genus  Denny  us 
Neumann.     Figures  63,  64.     Published  May  21,  1954  .    533-549 

New  species:  Dennyus  brevicapitis,  D.  intonsus,  D.  rotundocapitis, 
D.  brunneitorques,  D.  similis,  D.  spininotus,  D.  limbus. 

Cart  WRIGHT,  O.  L.  Scarabaeid  beetles  of  the  genus 
Bradycinettdus  and  closely  related  genera  in  the  United 
States.  Figures  14-16  and  plates  3,  4.  Published 
June  5,  1953 95-120 

New  genera:  Bolborhombus,  Bolbocerastes. 

New  species:  Bradycinetulus  rex,  Bolbocerastes  regalis,  B.  imperi- 

alis,  Bolborhombus  parvulus. 
New  subspecies:  Bolbocerastes  imperialis  kansanus. 

in 


IV  PROCEEDmGS    OF    THE    NATIONAL    MUSEUM 

Carvalho,  Jos6  C.  M.  Neotropical  Miridae,  LXIV: 
New  bugs  of  the  subfamily  Cylapinae  (Hemiptera). 
Figures  72-76  and  plate  15.     Published  March  2,  1955  .    621-632 

New  genera:  Brachyfulvhis,  Peritropoides. 

New    species:    Brachyfulvius    chapmi,   Peritroipoides   annulatus, 

P.    quadrinotatus,    Vannius    oculatus,    Fulvms    albonotatus,    F. 

castaneoiis,  F.  ornatifrons. 

GuRNEY,  Ashley  B.  Notes  on  the  biology  and  immature 
stages  of  a  cricket  parasite  of  the  genus  Rhopalosoma. 
Figures  8,  9  and  plate  1.     Published  March  10,  1953  .    .        19-34 

.  Distribution,  general  bionomics,  and  recogni- 
tion characters  of  two  cocla'oaches  recently  established 
in  the  United  States.  Figure  10  and  plate  2.  Pub- 
lished March  10,  1953 39-56 

Hershkovitz,  Philip.  Mammals  of  northern  Colombia, 
preliminary  report  No.  7:  Tapirs  (genus  Tapirus),  with 
a  systematic  review  of  American  species.  Figures  58- 
62.     Published  May  18,  1954 465-496 

Hyman,  Libbie  H.  North  American  triclad  Turbellaria, 
XIII:  Three  new  cave  planarians.  Figures  65-68. 
Published  June  14,  1954 563-573 

New    species:    Phagocata    caver  nicola,     Sphalloplana    georgiana, 
Speophila  hoffmasteri. 

Kenk,  Koman.  The  fresh-water  triclads  (Turbellaria)  of 
Alaska.  Figures  21-25  and  plates  6-8.  Published 
June  5,  1953 163-186 

New  species:  Phagocata  nivea,  Polycelis  borealis,  Dendrocoelopsis 
piriformis,  D.  alaskensis. 

Lachner,  Ernest  A.  A  revision  of  the  goatfish  genus  Upe- 
neus  with  descriptions  of  two  new  species.     Plates  13,  14. 

Published  May  18,  1954 497-532 

New  species:   Upeneus  asymmetricus,  U.  oligospilus. 

La  Rivers,   Ira.     Two  new  naucorid  bugs  of  the  genus 

Ambrysus.     Figure  1.     Published  February  12,  1953  .    .  1-7 

New  species:  Ambrysus  thermarum,  A.  bispimis. 

Lawrence,  R.  F.     Two  new  scale-mite  parasites  of  lizards. 

Figures  2-7.     Published  March  10,  1953 9-18 

New  species:  Geckobia  keegani,  G.  philippinensis. 

McDermott,  Frank  A.     Photuris  hethaniensis,  a  new  1am- 

pyrid  firefly.     Published  February  26,  1953 35-37 

New  species:  Photuris  bethaniensis. 


CONTENTS  V 

Morrison,  J.  P.  E.     The  relationships  of  Old  and  New  ^"^^ 

World  melanians.     Plate  11.     Published  April  20,  1954  .    357-394 

Pettibone,  Marian  H.  Marine  polychaete  worms  from 
Point  Barrow,  Alaska,  with  additional  records  from  the 
North  Atlantic  and  North  Pacific.  Figures  26-39.  Pub- 
lished May  21,  1954 203-356 

ScHULTZ,  Leonard  P.  Review  of  the  Indo-Pacific  anemone 
fishes,  genus  Amphiprion,  with  descriptions  of  two  new 
species.     Plates  9,  10.     Published  July  3,  1953 187-201 

New  species:  Amphiprion  tricinctus  Schultz  and  Welander,  Am- 
phiprion mauritensis  Schultz. 

AND  Marshall,  N.  B.     A  review  of  the  labrid  fish 

genus  Wetmorella  with  descriptions  of  new  forms  from  the 
tropical  Indo-Pacific.  Figures  52-54  and  plate  12.  Pub- 
lished April  21,  1954 439-447 

New  species:  Wetmorella  ocellata,  W.  albofasciata,  W.  triocellata. 
New  subspecies;  Wetmorella  philippina  bifasciata. 

Sturtevant,  a.  H.  Nearctic  flies  of  the  family  Periscelidae 
(Diptera)  and  certain  Anthomyzidae  referred  to  the  family. 
Published  June  16,  1954 551-561 

New  species:  Periscelis  occidentalis,  Cyamops  imitata. 

White,  Theodore  E.  Preliminary  analysis  of  the  fossil 
vertebrates  of  the  Canyon  Ferry  Reservoir  area.  Figures 
40-51.     Published  May  28,  1954 395-438 

New  genus:  Kentrogomphios. 

New    species:  Kentrogomphios    strophensis,    Paradjidaumo    spo- 
kanensis,  Eumys  cricetodontoides,  E.  latidens,  E.  spokanensis. 

Wirth,  Willis  W.  Biting  midges  of  the  heleid  genus  Stilo- 
hezzia  in  North  America.  Figures  11,  12.  Published 
May  15,  1953 57-85 

New  species:  Stilobezzia   (Neosiilobezzia)  stonei,  S.  (S.)  beckae,  S. 
(S).  punctipes,  S.  (S.)  sybleae,  S.  (S.)  thomsenae. 

.     American  biting  midges  of  the  heleid  genus  Mono- 


helea.     Figures  18,  19.     Published  June  17,  1953  ....    135-154 

New  species:  Monohelea  (Monohelea)  lanei,  M.  (M.)  macfiei,  M. 
(M.)  texana,  M.  (M.)  ornata,  M.  (M.)  stonei,  M.  (M.)  guianae, 
M.  (M.)  johannseni. 


PROCEEDINGS   OF   THE   UNITED   STATES   NATIONAL   MUSEUM 


Issued    BMrlrvA- ScjSB     ^y  '^* 


SMITHSONIAN  INSTITUTION 

U.  S.  NATIONAL  MUSEUM 


Vol.103  Washington:  1953  No.  331  1 

TWO  NEW  NAUCORID  BUGS  OF  THE  GENUS  AMBRYSUS 

By  Ira  La  Rivers^ 

The  following  new  Ambrysi  were  discovered  after  the  manuscripts 
dealing  with  United  States  and  Mexican  species  had  been  completed, 
and  this  paper  is  offered  as  a  supplement  to  these  treatments.  Keys 
are  included  to  allow  the  incorporation  of  the  new  species  into  the 
overall  keys  for  the  two  regions. 

Family  Naucoridae 

Genus  Ambrysus  Stdl,  1862 

Ambrysus  thertnarum,  new  species 

Figure  1,  &,  c 

Description. — A  small,  pale  species,  nearly  approaching  Airibrysus 
fwnebris  La  Rivers,  1949,  as  the  smallest  species  in  the  genus,  and 
greatly  resembling  the  latter  in  general  conspectus;  size  7.0-8.0  mm. 
long  and  4.5-5.0  mm.  wide.  Dorsum  bicolored,  the  scutellum  and  an- 
terior portion  yellowish,  the  posterior  area  (hemelytra)  brownish. 
Venter  brownish,  contrasting  strongly  with  the  yellow-white  ap- 
pendages. 

Head  glistening,  yellow,  slightly  roughened  and  weakly  punctate, 
protuberant  before  eyes  and  with  the  faintest  suggestion  of  trunca- 
tion in  front.    Eyes  blackish,  slightly  raised  above  general  head  sur- 

*  Department  of  Biology,  University  of  Nevada,  Reno,  Nev. 

228422—52  1 


2  PROCEEDINGS   OF    THE    NATIONAL   MUSEUM  vol.  los 

face  when  viewed  posteriorly.  Head  ratios  are :  Total  length  to  width 
(including  eyes)  19 :  28  (68  percent) ;  anterior  distance  between  eyes  to 
posterior  distance  12: 17  (71  percent)  ;  anterior  distance  between  eyes 
to  inner  eye-margin  12 :  11 ;  posterior  distance  between  eyes  to  greatest 
length  of  head  posterior  to  this  line  17 : 6  ( 35  percent) . 

Pronotum  glistening,  densely,  shallowly  punctate;  color  yellow  to 
amber ;  lateral  edges  smoothly  rounded,  nonserrate,  occasionally  with 
some  weak  pilosity.  Curvature  14  percent  (average  25:4),  postero- 
lateral angles  well  rounded.  Dorsal  ratios  are:  Width  between  an- 
terior angles  to  greatest  pronotal  width  29:  51  (57  percent) ;  median 
length  to  greatest  width  17:  51  (33  percent) ;  width  between  anterior 
angles  to  distance  between  anterior  angle  and  posterior  baseline  of 
pronotum  29 :  25  (86  percent). 

Scutellum  yellow-amber,  shiny;  shagreened  with  dense,  shallow, 
white-rimmed  punctation ;  ratio  of  three  sides  30 :  24 :  24. 

Hemelytra  uniformly  brownish,  yellowing  medially  and  in  em- 
bolar  region;  shagreened  as  is  scutellum.  Embolium  about  normal 
for  the  genus,  length-to- width  ratio  29:9  (31  percent),  rarely  with 
some  sparse,  occasional  marginal  pilosity.  Hemelytra  moderately 
exposing  lateral  connexival  edges,  which  latter  have  some  prominent, 
short  yellow  pilosity.    Hemelytra  just  attaining  abdominal  apex. 

Venter  reddish  brown,  contrasting  with  the  light-colored  legs. 
Connexival  posterolateral  angles  weakly  acute-angulate,  gradually 
enlarging  posteriorly,  not  in  the  least  spinose;  connexival  edges 
smooth,  nonserrate,  moderately  pilose.  Female  subgenital  plate 
apex  simply  concave,  the  concavity  somewhat  angular  at  its  deepest 
point.  Male  genital  process  sharply  right-angulate,  tapering  to  apex, 
appearing  much  like  a  very  thin  foot,  curving  abruptly  outward. 

Leg  I  yellow-white,  darkening  to  light  amber  on  tibia-tarsus; 
ratio  of  length  to  greatest  width  of  femur  27: 16  (59  percent)  ;  com- 
bined tibia-tarsus,  when  closed,  just  attaining,  or  slightly  surpassing, 
adjacent  (proximal)  end  of  femur. 

Leg  II  yellow ;  ratio  of  femoral  length  to  median  width  of  ventral 
surface  25:5  (20  percent),  length  1.9  mm.;  tibia  with  distal  end 
ventrally  with  one  complete  transverse  terminal  row  of  reddish  spines 
set  solidly  across  tip  (remnants  of  the  second,  subterminal  row,  when 
complete  and  unbroken  by  use,  extend  about  two-thirds  of  the  width  of 
tibia),  ratio  of  length  to  width  §2:  2  (10  percent),  length  1.75  mm. 

Leg  III  yellow;  ratio  of  length  to  femoral  width  30:  5.5  (18  per- 
cent), length  2.1  mm. ;  distal  end  of  tibia  ventrally  with  the  same  type 
of  transverse  spination  as  in  mesotibia ;  ratio  of  length  to  width  37 :  2 
(5  percent),  length  3.0  mm. 

Tibia  has  the  characteristic  spinate  appearance  in  contrast  to  the 
smoother  micro-armed  femora,  with  the  long,  silky,  swimming  hairs 
on  metatibia  contrasting  with  the  less  hirsute  appearance  of  the 


TWO    NEW   NAUCORID    BUGS — LA    RIVERS 


mesotibia.  All  tarsi,  except  protarsi,  3 -segmented ;  the  first  seg- 
ment minute ;  the  last  two  long  and  slender,  terminating  in  two  mod- 
erately curved  claws. 

Type  and  paratopes.— VSKM.  60987. 

Type  locality. — New  Mexico,  Sierra  County,  Hot  Springs. 

Material  examined. — Holotype  (male),  allotype,  and  four  para- 
types,  collected  by  Loew,  September  11,  1874  (P.  R,  Uhler  collection), 
in  U.  S.  National  Museum. 

Remarks. — This  unique  little  species,  discovered  during  the  course 
of  checking  the  Ambrysi  in  the  National  collection,  is  probably  an 
offshoot,  as  far  as  present  material  can  indicate,  of  Aimbrysus  morTnon 
stock.  The  female  subgenital  plate  is  mormonoid  in  structure,  and 
the  variable  7normo7i-ty'pe  male  process  could  easily  have  given  rise 


Figure  1. — Diagnostic  characteristics  of  Ambrysus  bispinus  and  A.  thermarum:  a.  Apex 
of  subgenital  plate  of  A.  bispinus,  female  holotype  (visible  only  in  ventral  view),  X22; 
b,  apex  of  subgenital  plate  of  A.  thermarum,  female  allotype,  X22;  c,  genital  process  on 
caudal  edge  of  tergite  V  of  A.  thermarum,  male  holotype,  X22;  d,  bispinate  condition 
of  left  connexivum  of  abdominal  sternite  V  of  A.  bispinus,  female  holotype,  showing  (1) 
anterior  posterolateral  angle  and  (2)  posterior  laterocaudal  angle  (ventral  view,  with 
caudal  tip  of  abdomen  oriented  at  the  top  of  the  field),  X22. 

to  the  distinctive  male  process  of  A.  thermarum,  which  has  evolved 
toward  the  A.  woodhuryi  type  to  such  an  extent  that,  with  only  the 
male  to  examine,  it  would  have  been  considered  an  aberrant  A.  wood- 
huryi. At  present,  it  is  the  most  definitely  adapted  thermal  ecad  of 
the  A.  mormon  type,  but  as  nothing  is  known  of  the  structure  of  the 
type  locality,  it  is  impossible  to  say  whether  this  seems  to  be  due  to 
complete  isolation  or  not. 


4  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  loe 

The  following  modification  of  my  key  to  United  States  Ambrysi 
(La  Rivers,  1951)  will  satisfactorily  compare  A.  thermanmi  with  its 
closest  relatives : 

8  (7).  Female  subgenital  plate  simply,  broadly,  and  smoothly  concave  apically, 
the  lateral  bordering  angles  sharp;  concavity,  measured  in  terms  of 
depth  of  concavity  against  length  between  bordering  lateral  angles,  never 
less  than  20  percent ;  in  clean  material  embolium  nearly  unicolorous,  only 
very  faintly  lighter  exteriorly,  no  contrasting  colors  present. 

buenoi  Usinger,  1946 

Female  subgenital  plate,  if  simply  concave,  has  (a)  rounded  lateral  angles, 

or  (b)  if  lateral  angles  are  sharp,  then  depth  of  concavity  is  considerably 

less  than  20  percent;  in  clean  material  embolium  always  sharply  and 

contrastingly  transversely  bicolored,  anterior  two-thirds  light  yellow, 

posterior  one-third  blackish-brown 8A 

8A  (8).  Male  genital  process  narrowing  to  apex,  strongly,  abruptly  and  almost 
right-angularly  turned  outward,  giving  the  appearance  of  a  long, 
attenuated  foot  with  a  rather  sharp  heel ;  female  subgenital  plate 
simply  concave  at  apex,  somewhat  angularly  so. 

thermarum,  new  species 

Male  genital  process  not  definitely  narrowing  to  apex,  and  not  foot-  or 

boot-shaped,  although  curving  variably  outward ;  female  subgenital 

plate  simply  and  roundly  concave  at  apex__  mormon  Montandon,  1909 

Ambrysus  bispinus,  new  species 

Figure  1,  a,  d 

Description. — A  large,  darkly  mottled  species,  well  rounded  and  con- 
vex; size  12.0-14.0  mm.  long  and  7.0-9.0  mm.  wide.  Dorsum  not 
distinctly  bicolored  (i.  e.,  anterior  portion  lighter  than  posterior  area) , 
more  or  less  uniformly  mottled.  Venter  yellow  or  brownish,  uni- 
colorous and  lighter  than  dorsum. 

Head  glistening,  yellow-brown,  smooth,  weakly  and  vaguely  punc- 
tate, slightly  and  broadly  protuberant,  and  truncate  before  the  eyes. 
Eyes  blackish,  nearly  flush  with  plane  surface  of  head  when  viewed 
posteriorly.  Head  ratios  are :  Total  length  to  width  (including  eyes) 
34:54  (63  percent)  ;  anterior  distance  between  eyes  to  posterior  dis- 
tance 26 :  33  (79  percent)  ;  anterior  distance  between  eyes  to  inner  eye- 
margin  26 :  25 ;  posterior  distance  between  eyes  to  greatest  length  of 
head  posterior  to  this  line,  33 :  8  (22  percent) . 

Pronotum  glistening,  yellowish  with  brown  mottling,  sparsely 
punctulate,  rugulose  behind  head;  lateral  edges  smooth,  nonserrate, 
rounded,  on  one  specimen  with  traces  of  faint  pilosity  which  appears 
to  have  been  rubbed  off.  Curvature  10  percent  (average  52:5); 
posterolateral  angles  well  rounded.  Dorsal  ratios  are:  Width  be- 
tween anterior  angles  to  greatest  pronotal  width  53 :  73  (73  percent) ; 
median  length  to  greatest  width  39 :  73  (53  percent)  ;  width  between 
anterior  angles  to  distance  between  anterior  angle  and  posterior  base- 
line of  pronotum  63 :  52. 


TWO    NEW    NAUCORID    BUGS — LA    RIVERS  5 

Scutellum  glistening,  yellow  medially  and  on  posterior  angle, 
yellow  along  lateral  edges,  reddish  brown  elsewhere;  shagreened  with 
dense,  coarse,  white-spotted  punctures ;  ratio  of  three  sides  70 :  50 :  50. 

Hemelytra  mottled  brown  and  yellow,  glistening ;  shagreened  as  is 
scutellum.  Embolium  slightly  narrow  for  the  genus,  length-to-width 
ratio  60: 18  (30  percent),  with  no  detectable  marginal  pilosity  in  the 
specimens  examined.  Hemelytra  moderately  exposing  connexival 
margins,  which  latter  have  sparse,  short  pilosity,  concentrated  some- 
what at  the  posterolateral  angles.  Hemelytra  just,  to  not  quite,  at- 
taining abdominal  tip. 

Venter  yellow  to  brown,  the  legs  usually  not  markedly  contrasting 
with  the  body  in  color.  Connexival  posterolateral  angles  acute-ang- 
ulate,  short,  gradually  increasing  in  size  posteriorly,  not  in  the  least 
spinose ;  margins  with  some  weak  dentation,  detectable  only  with  con- 
siderable magnification,  on  all  segments  except  I  and  II.  Female 
segment  V  (the  insect  being  oriented  so  that  the  abdominal  tip  is  at 
the  top  of  the  field)  possesses  two  posterolateral  connexival  processes ; 
the  exterior  posterolateral  angle  and  an  interior,  prominent,  blacker, 
rather  blunt  elongation  of  the  caudal  margin  of  segment  V  lying 
adjacent  to  the  posterolateral  angle.  I  have  referred  to  this  in  other 
species  as  the  laterocaudal  coimexival  angle.  Tip  of  female  sub- 
genital  plate  strongly  and  distinctively  quadrisinuate,  the  lateral  ter- 
minal angles  long  and  rather  sharp,  but  not  extending  as  far  caudad 
as  the  median,  paired  sinuosities  which  lie  close  together  as  a  doubly 
rounded,  blunt  process  longer  than  the  lateral  spines.  Character  of 
the  male  genital  process  unknown. 

Leg  I  amber ;  ratio  of  length  to  greatest  femoral  width  48 :  31 
(65  percent)  :  combined  tibia-tarsus,  when  closed,  just  attaining  adja- 
cent (proximal)  end  of  femur. 

Leg  II  amber;  ratio  of  length  to  median  width  of  femur  46:10 
(22  percent),  length  3.4  mm.;  tibia  with  distal  end  ventrally  with 
one-and-one-half  transverse  rows  of  reddish  spines,  the  terminal  row 
complete,  the  subterminal  row,  as  indicated  above,  only  half  complete : 
ratio  of  length  to  width  40 :4.5  (11  percent),  length  3  mm. 

Leg  III  amber;  ratio  of  length  to  median  femoral  width  57: 11  (19 
percent) ,  length  4.2  mm. ;  tibia  with  same  type  of  transverse  terminal 
spination  as  in  mesotibia,  ratio  of  length  to  width  70:  5  (7  percent), 
length  5.2  mm. 

Tibia  possesses  the  typical  spinate  appearance  in  contrast  to  the 
smooth  femora ;  long,  golden,  silky,  swimming  hairs  are  present  on  the 
metatibia  and  lacking  on  the  mesotibia.  All  tarsi,  except  protarsi,  are 
3-segmented,  the  first  segment  minute,  the  last  two  long  and  slender, 
terminating  in  two  moderately  curved  claws. 

Type  and  paratype. — USNM  60988. 


6  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  los 

Type  locality. — ^Mexico,  Veracruz,  Jalapa. 

Material  examined. — ^Holotype  (female)  and  a  single  female  para- 
type  from  Mexico,  Oaxaca,  Camotlan,  collected  by  W.  S.  Miller,  April 
14, 1944,  in  the  U.  S.  National  Museum. 

Remarks. — In  external  appearance,  A.  bispinus  is  superficially  in- 
distinguishable from  A.  cosmius  and,  as  such,  has  probably  shared 
the  latter's  confusion  as  A.  signoreti  Stal,  1862,  in  collections.  How- 
ever, the  structure  of  the  female  subgenital-plate  apices  is  quite  dif- 
ferent in  the  two  species.  In  this  respect,  A.  hispinus  most  closely 
resembles  A.  guttatipennis  Stal,  1876,  but  while  the  subgenital  plates 
might  not  be  able,  alone,  to  separate  all  variables  of  both  species,  the 
possession  of  primary  and  secondary  connexival  angles  on  abdominal 
segment  V,  with  present  material,  sharply  sets  A.  hispinus  off  from 
all  other  Ambrysi  of  the  signoreti  group.  These  two  angles  are  more 
specifically  referred  to  as  the  posterolateral  and  laterocaudal  con- 
nexival angles,  and  the  laterocaudal  is  lacking  in  all  other  known 
members  of  the  group.  It  has  shown  up  in  other,  distant  sections 
of  the  genus,  being  known  in  A.  parviceps  Montandon,  1897,  A.  hun- 
gerfordi  Usinger,  1946  (both  in  the  pudicus  group),  and  A.  puncti- 
collis  Stal,  1876  (a  monotypic  group).  It  is  this  character  upon 
which  the  specific  name  of  A.  hispinus  is  based.  In  addition,  the 
male  genital  process,  when  known,  will  furnish  more  evidence  of  the 
species'  affiliations,  either  supplementing  or  diverting  present  indica- 
tions. 

The  following  couplets  will  enable  the  new  species  to  be  incorpo- 
rated in  my  key  to  Mexican  Ambrysi  now  in  press : 

24  (15).  Female  subgenital  plate,  viewed  ventrally,  asymmetrical  on  left  side, 

where  a  prominent  flap  occupies  the  border  beyond  (laterad  of)  the 
left  lateral  angle ;  right  border  scarcely  produced. 

dilatus  Montandon,  1910 

Female  subgenital  plate,  viewed  ventrally,  not  markedly  or  noticeably 

asymmetrical  from  one  lateral  border  to  the  other 25 

25  (24).  Female  possessing  both  primary  and  secondary  posterolateral  connexi- 

val spines  on  abdominal  sternite  V  (i.  e.,  the  primary  angle  being  the 
posterolatei-al  angle  proper,  the  secondary  angle  being  the  laterocaudal 

angle) bispinus,  new  species 

Female  lacking  the  secondary  (laterocaudal)  angle. 

remaining  Mexican  species  of  the  signoreti  group 


TWO   NEW   NAUCORID    BUGS — LA   RIVERS  7 

References 

La  RrvEBS,  Ira 

1949.  A  new  species  of  Ambrysus  from  Death  Valley,  with  notes  on  the 

genus    in    the    United    States     (Hemiptera:    Naucoridae).     Bull. 
Southern  California  Acad.  Sci.,  vol.  47  (1948),  No.  3,  pp.  103-110. 

1950.  The  meeting  point  of  Ambrysus  and  Pelocoris  in  Nevada.     Pan-Pacific 

Ent.  vol.  26,  No.  1,  pp.  19-21. 

1951.  A  revision  of  the  genus  Ambrysus  in  the  United  States  (Hemiptera : 

Naucoridae).     Univ.  California  Publ.  Ent.,  vol.  8,  No.  7,  pp.  277- 
338,  illustr. 
—       The  Ambrysus  of  Mexico  (Hemiptera:  Naucoridae).    Univ.  Kansas 
Sci.  Bull.     (In  press.) 

MONTANDON,  A.  L. 

1897.  Hemiptera   cryptocerata. — Sous-fam.  Cryptocrieinae.     Verh.  zool.-bot. 

Ges.  Wien,  vol.  47,  pp.  6-23. 
1909.     Naucoridae.     Descriptions  d'especes  nouvelles.     Bull.  Soc.  Roum.  Sci., 

Bucharest,  vol.  18,  No.  1,  pp.  43-61. 
StAl,  Carl. 

1876.     Enumeratio  Hemipterorum.     Svenska  Vetensk.  Akad.,  vol.  14,  pt.  5, 

No.  4. 
UsiNGER,  Robert  L. 

1946.     Notes  and  descriptions  of  Ambrysus  St&l  with  an  account  of  the  life 

history  of  Ambrysus  mormon  Montd.     (Hemiptera:  Naucoridae). 

Bull.  Univ.  Kansas,  Sci.  Bull.,  vol.  31,  No.  1,  pp.  185-210. 


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PROCEEDINGS   OF   THE   UNITED   STATES   NATIONAL   MUSEUM 


SMITHSONIAN  INSTITUTION 
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Vol.   103  Washington:  1953  No.  3312 


TWO  NEW  SCALE-MITE  PARASITES  OF  LIZARDS 


By  R.  F.  Lawrence  ' 


Dr.  G.  W.  Wharton,  of  Duke  University,  Durham,  N.  C,  has  kindly 
submitted  to  me  for  identification  a  number  of  scale  mites  from  lizards 
of  the  families  Gekkonidae  and  Iguanidae.  The  material,  from  the 
Philippine  Islands,  consists  of  one  series  taken  from  a  lizard  at  Fort 
McKinley,  Rizal,  and  another  from  the  gecko  Hemidactylus  freTiatus 
in  a  house  at  Manila.  Both  series  consist  of  two  quite  different  species 
of  Geckobia.  These  are  described  in  this  paper  as  new  and  both  of 
them  have  been  taken  at  the  two  localities  mentioned  above.  There 
is  thus  good  reason  for  assuming  that  the  hosts  of  the  two  series  are 
the  same  and  that  the  "lizard"  of  Fort  McKinley  is  identical  with 
the  gecko  H.  frenatus  from  which  scale  mites  were  taken  at  Manila. 
All  these  parasites  were  collected  by  Dr.  H.  L.  Keegan,  Third  General 
Medical  Laboratory,  at  Fort  McKinley,  Rizal,  Philippine  Islands. 

A  further  series,  consisting  of  a  large  number  of  adults  and  larvae, 
were  collected  from  the  iguanid  lizard  Sceloporus  undulatus^  at  Dur- 
ham, N.  C,  by  Dr.  A.  S.  Pearse;  these  have  been  identified  as  Banks' 
New  World  pterygosomid  Geckohiella  texana. 

Several  species  of  the  scale  mite  parasite  Geckobia  may  live  on  the 
same  host;  Hirst  (1925,  p.  173)  has  already  pointed  this  out,  saying, 
"When  more  than  one  form  of  Geckobia  is  found  on  the  same  host, 
one  lives  beneath  the  ventral  scales  and  is  flattened,  being  considerably 
wider  than  long  and  having  scales  instead  of  hairs  on  the  venter. 
The  second  form  occurring  on  the  same  host  is  usually  to  be  discovered 
between  the  claw  and  pad  of  the  toes,  between  the  laminae  of  the  pad, 
or  between  the  toes  themselves;  this  form  is  practically  spherical  in 
shape  and  has  hairs  instead  of  scales  on  the  venter."    Although  there 

*  Natal  Museum,  Pietermaritzburg,  Union  of  South  Africa. 

228383—53  9 


10 


PROCEEDINGS    OF    THE    NATIONAL    MUSEUM 


is  no  definitive  information  that  the  two  strikingly  different  species 
of  Geckobia  here  described  actually  illustrate  Hirst's  observation,  it 
will  probably  be  found  that  the  two  forms  live  on  different  parts  of 
the  body  of  the  same  host. 

Family  Pterygosomidae 
Genus  Geckobia  Megnin,  1878 

Geckobia  keegani,  new  species 

Figures  2,  3 

Material  examined. — Three  females,  cotypes,  from  a  lizard,  Fort 
McKinley,  Kizal,  Philippine  Islands,  collected  by  H.  L.  Keegan,  Octo- 
ber 1948,  USNM  1931 ;  also  nine  females  bearing  same  locality  data 


Figure  2. — Geckobia  keegani,  new  species,  dorsal  surface. 


SCALE-MITE    PARASITES — LAWRENCE 


11 


Figure  3. — Geckohia  keegani,  new  species:  a,  Dorsal  scute;  h,  peripheral   dorsal   hair;   c, 
anterior  ventral  hair;  d,  posterior  ventral  hair. 

as  the  cotypes,  and  five  females  from  Hemidactylus  frenatus^  from 
Manila,  Philippine  Islands. 

Body  almost  circular;  dorsal  surface  as  in  figure  2,  with  fairly 
long,  almost  parallel-sided  hairs  differing  very  little  in  length,  but 
posteriormost  slightly  longer  than  anteriormost.  Dorsal  scute  well 
defined,  with  about  20  hairs  differing  very  little  in  general  appear- 
ance and  size  from  the  remaining  dorsal  hairs,  though  a  little  shorter 
than  the  longest  of  these. 

Ventral  surface  with  a  density  of  hairs  similar  to  that  of  the  dorsal 
surface  but  reaching  further  back,  almost  to  the  anal  field;  the  dis- 


12  PROCEEDINGS    OF    THE    NATIONAL    MUSEUM  vol.  io3 

crepancy  in  the  length  of  these  hairs  far  greater  than  in  those  of  the 
dorsal  surface,  anteriormost  a  little  less  than  half  as  long  as  posterior- 
most,  latter  a  little  shorter  than  the  longest  dorsal  hairs. 

Mouth  parts  normal,  pedipalps  short  and  stout,  both  second  and  third 
segments  above  with  a  setif  orm  hair,  that  of  the  second  weakly  barbed 
and  only  a  little  stouter  than  that  of  the  third,  which  is  smooth. 

Legs  equal-sized,  rather  small  in  proportion  to  the  body.  Pos- 
terior common  coxa  differing  from  most  species  of  Gechobia  in  having 
two  spurlike  hairs  on  its  anterior  and  only  two,  one,  or  even  none 
on  its  posterior  margin ;  usually  only  three  or  four  hairs,  instead  of 
the  normal  five,  are  found  on  this  coxa  and  these  are  comparatively 
small  and  weak.  The  basal  segments  of  the  legs  without  spurlike 
hairs  ventrally;  but  the  second  segment  in  all  legs  with  a  rather 
long,  slender,  and  weakly  barbed  hair  above. 

Dijnensions. — Width  of  body  0.603  mm.,  length  (including  mouth 
parts),  0.632  mm. 

Remarks. — The  species  closely  resembles  Gechohia  simplex  Hirst, 
1926,  described  from  Hemidactylus  leschenaulti.,  Madras,  India,  in 
the  shape  of  the  hairs  of  the  dorsal  surface  and  in  the  form  of  the 
dorsal  scute.  It  differs  from  this  species  in  having  only  about  half 
the  number  of  hairs  on  the  dorsal  scute,  these  being  also  more  similar 
to  the  remaining  body  hairs  in  length  and  thickness  than  is  the  case 
in  G.  simplex.  It  further  differs  in  there  being  very  little  difference 
in  the  shape  of  the  dorsal  hairs  of  the  second  and  third  palpal 
segments. 

Geckobia  philippinensis,  new  species 

Figures  4,  5 

Material  examined. — Two  females,  cotypes,  from  a  lizard.  Fort 
McKinley,  Rizal,  Philippine  Islands,  collected  by  H.  L.  Keegan, 
October  1948,  USNM  1932. 

Body  much  wider  than  long,  dorsal  surface  as  in  figure  4,  with  9  or 
10  large,  wide  hairs  in  the  area  usually  occupied  by  the  dorsal  scute, 
those  in  the  middle  of  the  body  much  smaller,  while  towards  the  pe- 
riphery posteriorly  and  laterally,  the  hairs  become  progressively  more 
elongate  and  pointed.  A  single  pair  of  centrally  situated  hairs  much 
longer  than  the  predominantly  short  hairs  by  which  they  are  sur- 
rounded.   Dorsal  scute  not  defined. 

Ventral  surface  as  in  figure  5,  a,  which  represents  a  section  in  the 
middle  line  of  the  body  extending  from  its  anterior  to  its  posterior 
margin;  the  transition  from  the  small  type  of  cylindrical  hair  to 
scales  of  lanceolate  form,  sudden  and  without  intermediate  types  of 
hair;  the  scales  toward  the  posterior  periphery  becoming  progres- 
sively more  elongate  and  narrow. 


SCALE-MITE    PARASITES — LAWRENCE 


13 


o 


14 


PROCEEDINGS    OF    THE    NATIONAL   MUSEUM 


Mouth  parts  normal,  pedipalps  short,  with  rounded  segments,  sec- 
ond segment  above  with  a  stout  plumose  hair  several  times  thicker 
than  that  of  the  third  segment,  which  is  smooth  and  setiform. 

Leg  IV  distinctly  longer  and  a  little  stouter  than  the  remaining 
ones;  posterior  common  coxa  as  in  figure  5,  e,  with  five  very  large 
and  distinct  conical   spurlike  hairs;   all  legs  with   a   fairly   stout 


d 


Figure  5. — Geckobia  philippinensis,  new  species:  a.  Hairs  of  ventral  surface  in  midline  of 
body;  b,  peripheral  dorsal  hair;  c,  small  hair  from  middle  of  dorsum,'^(/,  a  scalelike  ven- 
tral hair;  e,  posterior  common  coxa. 

plumose  hair  on  the  ventral  surface  of  segments  2  and  3  and  a 
long  setose  hair  on  the  dorsal  surface  of  segment  3,  that  of  leg 
IV  weakly  plumose. 

Additional  material  examined. — Two  females  from  the  same  host 
and  locality  as  the  types.  Dr.  Wliarton  sent  with  his  material  the 
lizard  host  from  which  the  Manila  specimens  were  taken.  From  the 
body  scales  of  this  gecko,  Hemidactylus  fre^iatus^  I  obtained  a  single 
female  of  the  species  described  above. 


SCALE -MITE    PARASITES — LAWRENCE  15 

Dimensions. — ^Width  of  body  0.400  mm.,  length  (including  mouth 
parts),  0.287  mm. 

According  to  Hirst's  key  (1925,  p.  174) ,  this  species  would  be  brack- 
eted with  Geckohia  australis  Hirst,  1926,  but  in  width  of  body  and 
general  form  of  dorsal  hairs  and  ventral  scales,  it  seems  to  resemble 
more  closely  G.  hindustanica  Hirst,  1926,  described  from  Hemidacty- 
lus  leschenaulti.,  Madras,  India.  It  differs  from  G.  hindustanica  in 
the  complete  absence  of  a  dorsal  scute  and  in  the  relatively  larger  size 
of  leg  IV,  as  well  as  in  other  details. 

Genus  Geckobiella  Hirst,  1917 

This  genus  was  erected  by  Hirst  (1917,  p.  138)  for  the  scale  mites 
of  Iguanid  lizards,  and  thus  far  its  members  have  been  found  to 
occur  only  on  species  of  the  genus  Sceloporus.  It  would  be  of  great 
interest  to  know  what  types  of  scale  mites  occur  on  the  other  numerous 
genera  of  the  family  Iguanidae  in  the  New  World. 

The  eyes,  which  are  not  mentioned  in  the  description  of  the  genus, 
consist  of  a  single  pair  situated  in  much  the  same  position  as  those 
of  Geckobia  and  Zomorobia.,  Lawrence,  1935,  being  located  anteriorly 
near  the  lateral  margin  of  the  body.  They  are  small,  only  a  little 
larger  han  the  ringlike  sockets  of  the  dorsal  hairs  (fig.  6,  &). 

Geckobiella  texana  (Banks),  1905 

Figures  6,  7 

OecJcobia  texana  Banks,  Proc.  Ent.  Soe.  Washington,  vol,  8,  p.  134,  1905. 
OeckoMella  texana   (Banks),  Hirst,  Ann.  Mag.  Nat.  Hist,,  ser.  8,  vol.  19,  p. 
138,  1917, 

Material  examined. — Numerous  adults  and  larvae  of  this  species 
collected  by  A.  S.  Pearse  from  Sceloporus  undulatus  at  Durham, 
N.  C,  in  January  1950. 

Adult  female. — ^Hirst  (1925,  p.  200,  fig.  19)  figured  only  the  ventral 
surface  of  this  species,  and  the  opportunity  is  now  taken  to  give  an 
illustration  of  the  dorsal  surface  (fig.  6) .  The  dorsal  hairs  are  much 
more  numerous  in  some  specimens  than  in  others,  there  being  probably 
a  certain  amount  of  variation  in  this  respect. 

The  mouth  parts  are  longer  and  more  robust  than  in  most  genera 
of  Pterygosomidae ;  the  free  portion  of  the  peritremes  reach  to  the 
distal  end  of  the  second  palpal  segment;  the  second  palpal  segment 
with  a  fairly  long  slender  hair  of  equal  thickness  throughout,  not 
pointed  at  the  apex,  with  fine  barbs ;  dorsal  hair  of  the  third  palpal 
segment  longer,  pointed  apically  and  smooth  or  almost  so ;  fourth  pal- 
pal segment  with  a  single  seta  above,  claw  stout,  conical,  and  somewhat 
curved. 


16 


PROCEEDINGS    OF   THE    NATIONAL   MUSEUM 


Figure  6. — Geckobiella  texana  (Banks):  a,  Dorsal  surface  of  female;  b,  eye. 

Leg  I  only  a  little  longer  than  IV,  distinctly  stouter  than  the  re- 
maining legs ;  segments  2  and  3  of  all  legs  with  a  setif  orm  hair  above, 
finely  barbed. 

Adult  male.—Th^  single  specimen  which  appears  to  be  an  adult 
male  is  considerably  smaller  than  the  female  and  with  far  fewer  hairs 


SCALE -MITE    PARASITES — LAWRENCE 


17 


Figure  7. — Geckobiella  texana  (Banks):  a.  Dorsal  surface  of  larva;  h,  penis  (?). 

on  the  dorsal  surface;  otherwise  it  does  not  differ  from  the  female; 
the  chitinous  genital  structures  are  rather  ill-defined,  the  large  penis 
(?)  (fig.  7,5),  projecting  slightly  from  the  genital  aperture  at  the 
posterior  extremity  of  the  body. 

Larva. — Dorsal  surface  with  very  few  symmetrically  disposed  hairs, 
as  in  figure  7,  a.  Eyes  present  in  the  same  position  as  the  adult.  Ven- 
tral surface  entirely  without  hairs  or  setae  except  for  a  single  seta  at 
the  base  of  legs  I  and  III.  Mouth  parts  closely  resembling  those  of 
the  adult,  but  the  peritremes  much  shorter,  reaching  to  a  little  below 


18  PROCEEDINGS    OF   THE    NATIONAL    MUSEUM  vol.  los 

the  base  of  the  second  palpal  segment,  and  the  claw  of  the  palp  more 
slender.    Legs  shorter  than  in  the  adult. 

Dimensions. — Female :  Length  of  body  0.913  mm. ;  width,  0.487  mm. 
Male :  Length  of  body  0.652  mm. ;  width,  0.326  mm.  Larva :  Length 
of  body  0.740  mm. ;  width,  0.413  mm. 

References 

BttEST,  ARTHtm  Stanley 

1917.  On  some  new  mites  of  the  suborder  Prostigmata  living  on  lizards. 

Ann.  Mag.  Nat.  Hist.,  ser,  8,  vol.  19,  p.  136. 
1925.  On  parasitic  mites  of  the  suborder  Prostigmata   (Trombidioidea),  on 
lizards.     Journ.  Linnaean  Soe.  London,  vol.  36,  p.  173. 
Laweence,  R.  F. 

1935.  The  prostigmatic  mites  of  South  African  lizards.    Parasitology,  vol.  27, 
No.  1,  p.  6. 


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Vol.   103  Washington  :  1953  No.  3313 

NOTES  ON  THE  BIOLOGY  AND  IMMATURE  STAGES  OF  A 
CRICKET  PARASITE  OF  THE  GENUS  RHOPALOSOMA 


By  Ashley  B.  Gurney  ^ 


Introduction 


For  more  than  three-quarters  of  a  century  wasps  of  the  genus 
Rhopalosoma  Cresson,  1865,  have  attracted  the  interest  of  hyme- 
nopterists  because  characters  of  the  adult  have  left  doubt  regarding 
family  relationships.  Although  the  family  Rhopalosomatidae  ^  was 
proposed  for  the  genus  in  1896,  there  remains  uncertainty  as  to  what 
other  genera  are  to  be  included  in  the  family.^  Neither  is  it  clear 
which  of  the  related  families  is  most  closely  allied  to  the  Rhopalosoma- 
tidae. Biological  studies  of  Rhopalosoma  together  with  a  critical 
examination  of  the  immature  stages  may  be  helpful  in  clarifying  these 
matters.  Until  now  the  only  original  published  information  concern- 
ing the  biology  and  immature  stages  is  that  by  Hood  (1914) .  He  col- 
lected a  larva  which  was  reared  to  maturity  and  identified  as  R.  poeyi 
Cresson,  which  is  currently  referred  to  R.  nearcticum  Brues.  Hood's 
account  is  brief  and,  while  it  is  a  good  general  indication  of  the 
biology,  contains  no  detailed  morphological  descriptions.  The  recent 
capture  of  a  larva  of  Rhopalosoma  at  Falls  Church,  Va.,  has  given 

^  Bureau  of  Entomology  and  Plant  Quarantine,  Agricultural  Research  Administration, 
U.  S.  Department  of  Agriculture. 

="  Originally  given  as  Rhopalosomidae  by  Ashmead  (18&6).  An  adult  specimen  of  the 
genus  has  been  illustrated  by  Westwood  (1874,  pi.  24,  fig.  9).  The  species  shown  there  is 
Rhopalosoma  aenigmaticum  (Westwood)  of  Hispaniola. 

8  Readers  are  referred  to  Brues  and  Melander  (1932,  p.  499),  Brues  (1943),  and  Krom- 
bein  (1951). 


228353—53 


19 


20  PROCEEDINGS    OF    THE    NATIONAL    MUSEUM  vol.  io3 

impetus  to  the  gathering  of  biological  data,  and  the  material  at  my 
disposal  has  also  permitted  morphological  studies  to  be  made  on 
several  of  the  immature  stages. 

I  am  much  indebted  to  Dr.  Henry  K.  Townes,  of  the  North  Carolma 
State  College  (in  absentia),  for  lending  specimens  of  Rhojyalosoma 
and  for  supplying  collecting  records  and  giving  helpful  suggestions. 
My  colleagues  Dr.  B.  D.  Burks,  K.  V.  Krombein,  and  Dr.  W.  H. 
Anderson  have  also  advised  me  generously.  David  G.  Shappirio,  of 
Harvard  University,  and  Dr.  T.  H.  Hubbell,  of  the  University  of 
Michigan,  have  kindly  supplied  data  on  parasitized  crickets  preserved 
at  the  University  of  Michigan  Museum  of  Zoology. 

Biological  observations 

The  larva  found  at  Falls  Church  ^  was  in  the  fourth  instar,  attached 
to  the  left  side  of  an  adult  female  of  Hapithus  agitator  Uhler.  While 
1  was  beating  deciduous  underbrush  for  psocids,  on  September  30, 
1950,  the  cricket  was  knocked  into  a  collecting  umbrella.  The  left 
hind  leg  was  held  at  an  unnatural  angle  with  the  body  and  the  cricket 
was  rather  inactive  and  easily  picked  up.  Leaves  and  other  organic 
material  were  added  to  the  jar  in  which  the  insect  was  placed,  but  the 
cricket  was  not  seen  to  feed.  The  next  day  it  scarcely  changed  its 
position,  though  the  antennae  occasionally  moved.  Meanwhile,  the 
ventral  surface  of  the  parasite  remained  securely  attached  to  the 
cricket  and  no  movement  was  noticed.  A  7  p.m.  on  the  evening  of 
October  2  the  larva  was  beginning  to  shed  its  skin,  as  shown  by  the 
appearance  of  the  white  integument  of  the  fifth  instar  where  the 
fourth  instar  skin  had  split  lengthwise  along  the  middle  of  the 
dorsum.  The  cricket  had  then  lost  its  foothold  and  was  dying  on  its 
side,  though  its  antennae  still  moved. 

By  8 :  30  p.m.  the  larva  (which  was  entirely  white  except  for  traces 
of  pale  brown  about  the  mandible)  had  left  the  dead  cricket  and  had 
moved  an  inch  or  so  away.  The  larva  was  placed  in  a  pill  box  of  soil 
where  it  gradually  darkened  to  a  creamy  yellow  color.  On  the  day 
following  the  molt  the  larva  made  frequent  twisting  contractions  as 
if  trying  to  burrow.  The  head  was  directed  downward  most  of  the 
time  and  one  silk  strand  coming  from  the  mouth  was  noticed.  The 
larva  did  not  succeed  in  becoming  completely  covered  with  soil ;  dur- 
ing the  next  day  it  was  relatively  inactive,  with  the  head  uppermost 


*  Generic  Identification  of  the  parasite  is  primarily  based  on  the  very  close  agreement 
with  the  situation  reported  by  Hood  (1914).  It  may  eventually  be  shown  that  more  than 
one  nominal  form  of  Bhopalosoma  inhabits  the  United  States  and  that  some  specimens 
from  the  vicinity  of  Washington,  D.  C.,  are  distinct  from  R.  nearcticum,  the  type  locality 
of  which  is  Kissimee,  Fla. 


CRICKET    PARASITE — GURNEY  21 

and  showing  occasional  movements  of  the  mouth  parts.  By  the  morn- 
ing of  October  5,  it  was  decidedly  quiescent  and  evidently  dying,  and 
by  noon  it  had  died  and  so  was  preserved  in  alcohol  (pi.  1).  If 
several  inches  of  fine  sand  had  been  provided  instead  of  heavy  gar- 
den loam,  perhaps  the  insect  w^ould  have  entered  the  soil  and  suc- 
cessfully spun  a  cocoon. 

The  present  larva  was  attached  to  the  cricket  on  the  membrane 
between  the  terga  and  sterna  of  abdominal  segments  1  to  &.  After 
the  fifth  instar  had  left,  the  cricket  (pi.  1)  bore  three  exuviae 
visible  from  above,  representing  the  second,  third,  and  fourth  instars. 
In  each  the  head  was  directed  posteriorly  and  located  farther  posteri- 
orly on  the  cricket  than  that  of  the  previous  instar.  The  strongest 
single  point  of  larval  attachment  was  apparently  just  anterior  to  the 
parasite's  modified  anus ;  there  the  attachment  was  quite  secure.  The 
larval  body  was  noticeably  curled,  but  the  entire  ventral  surface  was 
closely  appressed  to  the  cricket.  The  exuviae  of  larval  instar  1  was 
situated  transversely  on  the  ventral  surface  of  the  cricket,  in  a  groove 
just  behind  the  left  hind  coxa.  The  head  end  of  the  skin  was  directed 
mesally  on  the  cricket  and  extended  almost  to  the  sternal  plate  of  the 
first  abdominal  segment,  while  the  posterior  end  was  beneath  the  ends 
of  succeeding  exuviae.  Remnants  of  the  eggshell  occurred  with  the 
exuviae  of  the  first  instar.  On  the  body  of  the  cricket,  in  part  be- 
neath the  cast  skins  and  also  directly  adjacent  to  them  and  extending 
briefly  onto  the  left  tegmen  (front  wing),  was  an  accumulation  of 
dried  mucus  or  similar  material.  This  was  scalelike  in  appearance 
and  I  first  mistook  it  for  exuviae.  The  larval  head  was  not  inserted 
through  the  cricket's  body  wall,  nor  was  there  any  definite  opening, 
but  the  membrane  where  the  last  feeding  mandibles  (fourth  instar) 
were  applied  showed  signs  of  having  been  pierced.  The  points  of 
contact  for  the  mandibles  of  the  second  and  third  instars  were  covered 
with  the  exuviae  of  later  instars. 

A  second  larva  of  RJiopalosom-a  in  the  U.  S.  National  Museum  was 
collected  on  a  last  instar  male  nymph  of  Hapithus  sp.  at  Marietta,  Ga., 
August  9,  1947.  The  larva  is  in  the  fifth  instar,  though  it  probably 
was  captured  in  an  earlier  stage  since  the  host  cricket  is  also  preserved. 
The  exuviae  which  were  attached  to  the  left  side  of  the  cricket  agree 
with  those  of  the  Falls  Church  specimen  so  far  as  has  been  noted. 

The  third  larA^a  of  Ehopalosoina  examined  (fig.  9,  e,  f)  was  preserved 
in  the  fourth  instar  after  having  been  collected  on  an  adult  female  of 
Hapithus  agitator  at  Fuguay,  N.  C,  September  12,  1950,  by  Townes, 
Rabb,  and  Howden  (North  Carolina  State  College  collection)  (pi.  1). 
The  head  of  the  final  feeding  stage  of  the  larva  was  on  the  cricket's 
fifth  abdominal  segment,  with  the  mandibles  applied  to  the  mem- 
brane  close   to   the   spiracle   and   nearer  the  tergum   than  to  the 


22  PROCEEDINGS    OF    THE    NATIONAL    MUSEUM  vol.  io3 

sternum.  A  crust  of  a  dried  secretion  was  present,  as  in  the  case  of 
the  Falls  Church  larva.  Though  a  crust  may  have  been  associated 
with  the  Marietta,  Ga.,  material,  none  was  seen.  The  eggshell  and 
first  larval  exuviae  of  each  of  the  three  larvae  occurred  immediately 
behind  the  hind  coxa  of  the  cricket.  Likewise,  the  hind  leg,  of  each 
cricket,  adjacent  to  the  parasite,  was  held  outward  as  shown  in  the 
photographs. 

A  fourth  larva  examined  is  one  of  several  which  Dr.  Townes  has 
told  me  of  finding  on  Hapithus  and  Orocharis.  Of  this  particular 
larva  he  has  written,  "On  September  12,  1950, 1  collected  (at  Fuguay, 
N.  C.)  a  large  Orocharis  nymph  with  a  one-third  grown  larva  on  it. 
I  kept  the  cricket  in  a  jar  and  its  parasite  prospered.  On  September 
17  I  found  the  remains  of  the  demolished  nymph  in  the  jar  and  the 
Rhopalosoma  larva,  now  creamy  white  with  a  large  head  and  pro- 
jecting mandibles,  busily  hitching  itself  along  with  its  mandibles. 
It  was  very  restless,  so  I  put  it  in  some  damp  sand.  The  larva 
promptly  worked  into  the  sand  and  came  to  rest  in  one  corner  of  the 
jar,  under  the  sand.  Three  days  later  it  had  completed  an  elongate, 
castaneous  cocoon." 

The  cocoon  was  kept  indoors  by  Dr.  Townes,  at  a  temperature  in 
the  neighborhood  of  55°  F.  In  late  February  he  mailed  it  to  me  and 
suggested  that  dissection  to  determine  whether  pupation  had  occurred 
might  be  advisable.  The  parchmentlike  cocoon  was  14  millimeters 
long,  about  5  millimeters  in  diameter,  and  rounded  at  the  ends.  The 
inner  surface  was  a  lighter  chestnut  color  than  the  outside  and  scarcely 
any  individual  threads  of  silk  were  apparent  except  on  the  outside. 
When  the  cocoon  was  opened  on  February  23,  the  larva  was  doubled 
up  in  the  middle  of  it  with  about  one-fourth  of  the  cocoon's  length 
empty  at  each  end.  Pupation  had  not  occurred,  nor  had  there  been 
any  casting  of  larval  skins  or  passing  of  fecal  material,  but  the  larva 
appeared  perfectly  healthy.  After  preservation  it  was  6  millimeters 
long.     The  mandibles  bore  no  teeth. 

The  original  observations  of  Hood  (1914)  were  based  on  a  larva 
taken  at  Plummer's  Island,  Md.,  October  6, 1912.  The  host,  Orocharis 
saltator  Uliler,  was  quite  active  when  found,  as  it  was  said  to  be 
"scampering  over  the  forest  floor  as  rapidly  as  a  large  abdominal 
protuberance  and  a  nearly  functionless  hind  leg  would  permit."  The 
anterior  end  of  the  parasite  was  at  the  host's  eighth  abdominal  seg- 
ment. Tliis  specimen  grew  rapidly,  left  the  cricket  three  days  after 
capture  and  entered  the  soil,  emerging  indoors  as  an  adult  about 
March  1,  1913.  The  information  Hood  gave  concerning  the  exuviae 
and  the  quiescent  state  of  the  host  just  prior  to  the  parasite's  final 
visible  molt  agrees  fully  with  my  findings.  The  cocoon  from  which  the 
adult  emerged  has  recently  been  opened  and  the  fifth  instar  exuviae 


CRICKET   PARASITE — GURNEY  23 

were  relaxed  and  studied.  The  mandibles  of  the  exuviae  differ  some- 
what in  shape  from  those  of  the  Falls  Church  larva  and  they  bear  no 
teeth. 

In  the  University  of  Michigan  Museum  of  Zoology  are  at  least  20 
pinned  cricket  specimens  bearing  parasitic  larvae  presumed  to  be  of 
one  or  more  species  of  Rhopalosoina.  I  have  not  examined  them, 
but  detailed  host  data  have  been  furnished  me,  together  with  the  state- 
ment that  the  larvae  vary  considerably  in  size  and  that  all  are  brown. 
The  specimens  were  collected  by  Dr.  T.  H.  Hubbell  and  several  asso- 
ciated collectors  and  former  students.  The  hosts,  with  the  number 
of  parasitized  specimens.  States  where  collected,  and  generalized 
dates  of  collection  are  as  follows :  Hapithus  agitator  agitator  Uhler, 
seven  specimens,  southeastern  Indiana,  central  Missouri,  Mississippi, 
Tennessee,  all  collected  in  the  last  half  of  August ;  H.  agitator  quad- 
ratus  Scudder,  four  specimens,  Florida,  late  July  to  early  September; 
H.  hrevipennis  Saussure,  four  specimens,  Florida,  late  July  to  mid- 
November  ;  Orocharis  saUator  Uhler,  four  specimens,  July  4  to  July 
29.  The  host  associated  with  an  additional  larva  from  Florida  is 
not  recorded.  These  records  represent  the  parasitism  of  this  type 
occurring  among  several  hundreds  of  Hapithus  and  Orocharis 
specimens. 

The  specimen  of  Hapithus  agitator  on  which  the  Falls  Church  larva 
was  parasitic  was  preserved  in  alcohol  and  later  dissected  to  determine 
the  condition  of  its  internal  organs.  Scarcely  any  traces  of  food 
were  in  the  digestive  tract,  none  of  recent  ingestion.  Keproductive 
organs  were  evidently  intact,  but  there  were  no  well-developed  eggs 
or  other  signs  of  normal  functioning  of  the  organs. 

The  impression  created  was  that  the  cricket  had  been  weakened 
to  a  point  where  the  drain  on  its  vitality  finally  caused  death,  though 
there  had  been  no  mechanical  destruction  of  organs. 

Various  details  of  the  biology  remain  to  be  clarified,  but  the  main 
outline  seems  evident.  Upon  hatching,  the  larva  probably  crawls  part 
way  from  the  eggshell  and  starts  its  development  in  a  more  or  less 
transverse  position.  At  the  first  molt  the  larva  apparently  shifts 
the  anterior  part  of  the  body  so  that  it  extends  longitudinally  along 
the  side  of  the  cricket,  though  keeping  the  same  place  of  anal  attach- 
ment. The  third  and  fourth  instars  each  move  the  head  posteriorly 
on  the  cricket  at  molting  time.  Sharp  mandibles  occur  in  the  second, 
third,  and  fourth  instars,  and  these  apparently  pierce  the  cricket's 
integument  and  permit  fluids  to  be  taken  into  the  mouth.  Definite 
mouth  parts  of  the  first  instar  have  not  been  seen,  but  may  occur. 
The  fifth  instar  does  not  feed  but  is  an  inactive  stage  that  leaves  the 
host  and  spins  a  cocoon  in  the  soil.  Pupation  is  evidently  delayed, 
perhaps  until  near  emergence  time  of  the  adult. 


24  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.103 

Regarding  habits  of  Rhopalosoma  adults,  Dr.  Townes  has  furnished 
the  following  notes :  "I  have  seen  the  adults  on  several  occasions  and 
usually  was  fortunate  to  catch  them  when  seen.  I  should  say  that 
in  the  Upper  and  Lower  Austral  Zones  of  the  Atlantic  States  they 
are  widespread  and  not  uncommon  in  the  right  habitat.  This  appears 
(.0  be  places  of  dense,  shrubby  vegetation  where  there  is  considerable 
humidity,  as  along  stream  bottoms  and  seashores,  or,  in  other  words, 
the  same  sorts  of  places  where  Hapithus  and  Orochuris  abound.  The 
adults  may  be  flushed  from  the  bushes.  They  fly  up  and  alight  again 
like  an  ophionine  ichneumonid.  They  may  be  distinguished  from 
ophionines  in  flight  only  by  a  slightly  stockier  appearance.  At  Long 
Beach,  North  Carolina,  at  twilight  on  July  9,  1949,  I  saw  about  10 
all  together  in  normal  flight,  cruising  in  a  rather  slow  but  erratic 
manner  just  over  the  beach  shrubbery,  again  like  an  ophionine  ich- 
neumonid in  its  twilight  flight.  They  were  flying  until  no  longer 
visible  in  the  gathering  dark.  I  could  catch  only  two  of  these  and 
they  were  both  males." 

The  following  collection  dates  accompanying  specimens  identified 
as  Rhopalosoma  nearcticum  are  available  as  an  indication  of  when 
adults  are  active :  Maryland :  Mayo  Beach,  September  14, 1947,  female 
(Townes);  Berlin,  July  19,  1932,  female.  North  Carolina:  Long 
Beach,  July  9,  1949,  two  males  (Townes)  ;  Wallace,  August  2,  1949, 
female  (Townes).  South  Carolina:  Greenville,  September  2,  1940, 
male  (H.  and  M.  Townes).  Georgia:  Stone  Mountain,  August  13, 
1949,  female  (Fattig)  ;  Atlanta,  August  11,  1946,  female.  Florida : 
About  a  dozen  specimens,  essentially  all  reported  by  Brues  (1943), 
ranging  from  June  9  to  August  13,  but  mostly  collected  in  June  and 
July. 

In  addition  to  the  foregoing  States,  nearcticum  has  been  recorded 
from  Missouri  and  Kentucky  by  Krombein  (1951). 

The  known  hosts  of  Rhopalosoma  in  the  United  States  are  Orocharis 
saltator  Uhler  and  Hapithus  agitator  Uhler.  Judging  from  the  rec- 
ords of  parasitized  crickets  preserved  in  the  University  of  Michigan 
Museum  of  Zoology,  Hapithus  h'revipennis  Saussure  appears  to  be  a 
host  also.  The  distribution  of  0.  saltator,  as  stated  b}^  Hebard  (1931, 
p.  217) ,  covers  the  entire  southeastern  United  States,  extending  north- 
ward to  southeastern  Pennsylvania,  Illinois,  and  Missouri,  and  un- 
doubtedly to  eastern  Texas  in  the  Southwest.  In  southern  Florida  a 
primarily  West  Indian  species,  Orocharis  gryUodes  (Pallas) ,  occurs. 
H.  agitator  occurs  as  far  northeast  as  Long  Island,  and  otherwise  is 
largely  comparable  to  O.  saltator  in  distribution  (Blatchley,  1920, 
p.  740;  Hebard,  1931,  1938;  Rehn  and  Hebard,  1916,  pp.  308-310). 
Two  geographic  subspecies  are  recognized,  the  southeastern  one,  H. 
agitator  quadratus  Scudder,  living  from  the  Atlantic  Coast  to  Browns- 


CRICKET    PARASITE — GURNEY  25 

ville,  Tex.,  and  in  the  East  apparently  merging  with  typical  H.  agita- 
tor in  central  Georgia  and  along  tlie  "fall  line"  in  North  Carolina.  H. 
hrevipennis  inhabits  Florida,  Georgia,  and  Louisiana,  and  doubtless 
other  southeastern  States.  It  may  be  that  any  of  the  crickets  men- 
tioned above  are  satisfactory  hosts  of  Rhopalosoma.  In  southeastern 
Pennsylvania  adults  of  both  Orocharis  and  Hapithus  begin  to  appear 
by  early  August,  and  in  all  of  the  more  northern  States  the  crickets 
pass  the  winter  as  eggs,  and  adults  persist  until  frost  time  in  the  fall. 
Blatchley  (1920)  reported  finding  hibernating  adults  of  E.  agitator 
in  Florida  and  suggested  that  they  represented  a  fall  generation.  The 
subfamily  Eneopterinae,  to  which  Eapithus  and  Orocharis  belong,  is 
primarily  tropical,  and  various  genera  are  well  represented  in  the 
Neotropical  Region,  from  which  Brues  (1943)  has  recognized  four 
species  of  Rhopalosoma. 

Morphological  descriptions 

Mature  larva  {fifth  instar  from  Falls  Churchy  Va.). — General  shape 
as  in  plate  1;  strongly  mandibulate;  legs  entirely  absent;  abdomi- 
nal segments  moderately  well  demarked,  less  so  posteriorly,  those  be- 
yond segment  8  not  readily  separated;  tAVO  thoracic  and  eight 
abdominal  pairs  of  spiracles. 

Head  in  frontal  view  as  in  figure  8,  / ;  occipital  border  broadly  and 
evenly  rounded  (asymmetrical  in  the  figure  because  it  is  bordered  by 
uneven  cervical  folds)  ;  vertex  and  frons  not  individually  demarked, 
the  integument  shiny,  slightly  irregular,  not  noticeably  punctate ;  no 
frontal  arms  evident;  paired  conspicuous  spots  (ey),  presumed  to  be 
vestigial  eyes,  just  above  base  of  clypeus ;  antenna  reduced  to  sensory 
pit  with  traces  of  facetlike  division,  situated  well  to  the  side  of  frons, 
no  apparent  projecting  sensilla;  a  presumed  sensory  area  represented 
by  about  nine  poorly  defined,  pigmented  spots  (sa),  with  tiny  seta  at 
lateral  margin  of  group;  frons  and  vertex  with  very  sparse,  short 
setae,  grouped  as  illustrated;  six  strong  clypeal  setae;  labrum  con- 
spicuously bilobed,  each  lobe  with  two  major  setae,  lateral  and  an- 
terior margins  with  microsetae,  an  irregiilar  row  of  pits  across  each 
lobe,  some  bearing  short  setae;  mandibles  heavily  sclerotized,  apex 
prominent  and  narrowly  rounded,  basal  third  swollen  laterally  and 
with  a  single  seta,  biting  margin  thin,  demarked  from  main  mandib- 
ular structure  by  groove,  and  sharply  but  delicately  toothed.  (Left 
mandible  with  four  well-spaced  teeth  along  middle  third  of  biting 
margin,  followed  by  closely  set  group  of  four;  right  mandible  with 
two  teeth,  others  perhaps  lost  by  breakage.)  Maxilla  with  several 
medium-sized  setae  as  illustrated,  apex  near  tip  of  mandible  sharply 
rounded  and  with  asperities,  maxillary  palpus  barrel  shaped,  galea 


26  PROCEEDINGS    OF    THE    NATIONAL    MUSEUM  vol.  io3 

equally  elongate  but  of  smaller  diameter ;  labium  with  palpi  subequal 
to  galeae,  area  of  sericteries  with  two  angular  projections  posterior  to 
weakly  sclerotized  nonsegmented  elongate  appendages  of  uncertain 
homology;  apparent  opening  of  silk  duct  (sd)  between  bases  of  latter 
appendages. 

Anterior  spiracle  on  posterior  third  of  prothorax;  second  spiracle 
barely  anterior  to  groove  separating  meso-  and  metathorax;  abdominal 
spiracles  on  segments  1  to  8,  slightly  more  dorsad  than  thoracic  ones, 
situated  near  anterior  margin  of  the  respective  segments ;  each  spiracle 
unspecialized  so  far  as  visible  under  72-power  magnification,  consist- 
ing of  circular  disk  with  central  aperture  (internal  structures  not 
studied)  ;  no  setae  on  thorax  or  abdomen  apparent;  no  apparent  spe- 
cializations for  crawling  or  clinging;  vestigial  anus  transverse,  at 
posterior  margin  of  segment  10. 

Coloration  :  General  color  creamy  yellow,  darker  on  head  (con- 
spicuous black  areas  on  body  shown  in  photograph  are  necrotic  spots 
developed  at  time  of  death).  Vertex,  frons,  and  clypeus  yellowish; 
eye  spots,  sensory  areas,  antennae,  and  pits  at  bases  of  setae  pale  brown- 
ish ;  a  darkened  area  anterior  to  sensory  area ;  a  somewhat  darkened 
indefinite  band  across  base  of  clypeus  connecting  bases  of  mandibles ; 
labrum  colorless,  except  for  brown  setae  and  pits ;  mandibles  marked 
as  illustrated  with  rich,  dark  brown ;  maxillary  and  labial  palpi  and 
galeae  dark  brown;  labium  white,  elongate  appendages  in  area  of 
sericteries  tinged  with  yellowish ;  spiracles  yellowish ;  no  special  pig- 
mented areas  about  nonfunctional  anus  or  elsewhere  on  thorax  and 
abdomen. 

Measurements  (in  millimeters)  :  Length  8  (would  actually  be 
slightly  longer  if  body  not  curved);  greatest  width  of  body  3.2; 
greatest  width  of  head  capsule  1.3. 

Comments:  Fifth  instar  from  Marietta,  Ga.,  more  robust  than 
larva  from  Falls  Church,  Va. ;  setae  on  frons  and  vertex  smaller, 
scarcely  noticeable;  teeth  on  mandibles  lacking.  Measurements  (in 
millimeters)  :  Body  length  8.5 ;  greatest  width  4 ;  greatest  width  of 
head  capsule  1.6. 

Penultmiate  stage  larva  {fourth  instar  from  Fuguay^  N .  C). — Gen- 
eral appearance  typically  hymenopterif  orm  (fig.  9,  e,  /)  ;  head  capsule 
and  mouth  parts  not  so  highly  developed  as  in  fifth  instar ;  legless ;  10 
pairs  of  spiracles ;  modified  anus  apparently  specialized  as  a  hold-fast 
organ. 

Head  capsule  (fig.  8,  c)  rather  masklike  in  contrast  with  that  of 
fifth  instar,  integument  very  rough  and  irregularly  rugose,  a  conspicu- 
ous tranverse  ridge  (epistoma)  dorsad  of  labrum;  two  pairs  of  blunt 
protuberances  on  upper  part  of  face,  the  median  pair  (mt)  broad  and 


U.  S.  NATIONAL  MUSEUM 


PROCEEDINGS.  VOL.  103  PLATE  1 


u  = 


£1 


5=< 


V   a; 

•-2  ~ 


5p 


DO 


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P 


CRICKET   PARASITE — GURNET 


27 


labs 


mxs 


FiQUEE  8.— The  genus  Rhopalosoma  Oresson,  1865:  o,  Mouthparts  of  fourth  Instar  exuviae;  6,  external  view 
of  sixth  abdominal  spiracle  of  fourth  instar  exuviae;  c,  front  view  of  head,  fourth  instar  alcoholic  specimen; 
d,  antenna  of  fourth  instar  exuviae;  e,  anal  region,  fourth  instar  exuviae;  /,  front  view  of  head,  fifth  instar 
alcoholic  specimen;  (c  made  from  Fuguay,  N.  O.,  specimen,  all  others  from  Falls  Church,  Va.,  material). 


28  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  108 

irregularly  rounded,  lateral  pair  (It)  narrow  and  evenly  rounded;  an- 
tennal  socket  (an,  fig.  9,  e)  circular,  minute,  traces  of  sensilla  scarcely 
visible  under  72-power  magnification;  labrum  membranous  except 
mesally  and  at  lateral  extremities,  strongly  bilobed,  each  lobe  bearing 
two  small  setae  about  equidistant  from  center  and  a  minute  seta  near 
anteromesal  margin ;  exposed  apical  fourth  of  overlapping  mandibles 
triangular,  the  apex  sharply  acute,  about  seven  sharp  teeth ;  approxi- 
mate basal  third  of  each  mandible  exposed  near  end  of  epistoma,  bear- 
ing a  single,  minute  seta ;  a  long  tapering  apodeme  extending  dorso- 
laterally  within  head  capsule;  maxillary  sclerome  (mxs)  anterior  to 
base  of  mandible,  shaped  as  illustrated ;  labial  sclerome  (labs)  an  ir- 
regular, incomplete  circle. 

Abdomen  with  10  apparent  segments;  spiracles  located  as  in  fifth 
instar ;  entire  dorsal  area  glabrous ;  venter  of  prothorax  and  cervical 
region  bearing  minute  asperities ;  dorsal  and  lateral  integument  ap- 
pearing as  if  covered  with  closely  appressed  scales  (see  later  descrip- 
tion of  exuviae)  ;  eight  abdominal  segments  with  ventral,  transverse 
groups  of  small  platelets ;  hold-fast  organ  at  posterior  margin  of  seg- 
ment 10,  transverse,  weakly  crescent-shaped,  with  short  membranous 
flap  extending  posteriorly. 

Coloration:  General  color  dark  blackish-brown;  ventral  region 
(unstippled  areas  on  fig.  9,  e,  /)  unpigmented;  a  white  middorsal  line 
(where  split  occurs  at  subsequent  molt)  extending  entire  length  of 
body ;  head  considerably  darker  than  thorax  or  abdomen ;  region  of 
frons  lighter  colored  than  main  epicranial  region,  two  darker  pig- 
mented spots  (ps)  anterior  to  median  tubercles,  a  wide  longitudinal 
white  stripe  on  frons,  interrupted  on  anterior  portion  of  vertex  and 
resumed  as  narrow  line  on  occiput;  labrum  unpigmented  except  at 
lateral  extremities,  on  a  slender  transverse  basal  median  sclerite,  and 
mesally  at  the  junction  of  the  lobes;  mandibles  and  scleromes  of  max- 
illae and  labium  pigmented  with  brown  as  illustrated.  Abdomen 
with  paired  unpigmented  spots  dorsad  of  spiracles  3  to  8,  and  in  a 
similar  position  on  abdominal  segment  9  (which  bears  no  spiracle) ; 
transversely  grouped  platelets  of  ventral  surface  pale  brown,  also  clos- 
ing sclerites  of  hold-fast  organ  and  immediately  adjacent  membrane. 

Measurements  (in  millimeters)  :  Length  of  body  5.7;  greatest 
width  of  body  2.2 ;  greatest  width  of  head  capsule  1.4. 

Comments:  Slide-mounted  preparations  of  the  cast  skin  of  the 
fourth  instar  from  Falls  Church,  Va.,  permitted  structures  to  be 
observed  in  much  greater  magnification  than  was  possible  with  the 
preserved  alcoholic  larva  from  Fuguay,  N.  C.  While  no  significant 
differences  in  these  individual  fourth  stage  larvae  apparently  occur, 
it  is  important  that  the  features  illustrated  and  described  be  attributed 
to  the  correct  specimens. 


CRICKET   PARASITE GURNEY 


29 


d  cc      <:^o©'^^^20 


'^^::>z>   c^ 


hg--^ 


Figure  9— The  genus  Rhopalosoma  Cresson,  1865:  a.  Section  from  lateral  area  of  thorax,  showing  closely 
grouped  platelet  structure,  fourth  instar  exuviae;  6,  integumental  asperities  on  ventral  surface  of  pro- 
thorax,  fourth  instar  exuviae;  c,  d,  specialized  platelets  on  ventral  surface  of  fourth  and  fifth  abdominal 
segments,  respectively,  fourth  instar  exuviae;  e,  f,  ventral  and  lateral  views,  respectively,  of  fourth  instar 
alcoholic  specimen;  (e,f,  from  Fuguay,  N.  C,  specimen,  others  from  Falls  Church,  Va.,  material). 


an— antenna 
cv — closing  valves 
ep — epistoma 
es— esophagus 
ey— eye 
ga—  galea 
hg— hind  gut 


labs— labial  sclerome 
labr — labrum 
Ip— labial  palpus 
It— lateral  tubercle 
md — mandible 
mdc — mandibular  condyle 
mt— median  tubercle 


mxp— maxillary  palpus 
mxs — maxillary  scleroma 

ps— pigment  spots 

sa — sensory  area 

sd— opening   of  silk    gland 
or  sericteries 

tr— trachael  ring 

vp— ventral  platelets 


30  PROCEEDINGS    OF    THE    NATIONAL    MUSEUM  vol.  io3 

Exuviae  split  lengthwise  on  dorsum,  from  epistoma  to  tenth  ab- 
dominal segment;  presumed  antenna  consisting  of  circular  socket 
(jBg.  8,  d)  with  three  tiny  unpigmented  circles  in  center,  each  bearing 
a  central  spot  of  pigment;  mouth  parts  (fig.  8,  a)  with  mandibles 
the  most  characteristic  and  best  preserved  of  the  parts ;  mandible  thin, 
much  flattened,  biting  margin  with  large  apical  tooth  and  about  10 
acute  smaller  teeth  in  a  single  row  along  apical  third,  an  emargination 
a  little  posterior  to  the  teeth,  portion  of  margin  overlaid  by  labrum 
scarcely  pigmented,  a  well-sclerotized  and  pigmented  region  at  base 
of  mesal  margin;  posterior  margins  of  mandible  with  a  prominent 
black  ventral  condyle,  a  small  seta  near  base  of  long  lateral  apodeme 
(presumed  mesal  apodeme  for  muscle  attachment  missing  in  available 
material)  ;  labrum  largely  membranous,  with  irregular  transverse 
wrinkles,  a  constricted  pigmented  sclerite  mesally,  about  five  subapical 
minute  pits  arranged  transversely  on  each  lateral  half;  a  poorly 
defined,  somewhat  striated  organ  leading  posteriorly  to  labial  sclerome 
from  base  of  labrum,  dorsad  of  remnants  of  apparent  esophagus 
(es) ;  labial  and  maxillary  scleromes  showing  somewhat  different 
shape  in  flattened  aspect  than  in  natural  position  on  alcoholic  larva. 

Integument  of  most  of  thorax  and  abdomen  smooth,  microscopically 
marked  into  platelets  of  different  shapes  on  various  areas  of  the 
body,  averaging  transversely  elongate  ventrally,  irregularly  quadrate 
(fig.  9,  a)  dorsally  on  thorax  and  anterior  half  of  abdomen;  ventral 
region  of  neck  and  prothorax  bearing  minute  asperities  of  varied 
character  (fig.  9,  J) ,  some  setiform,  others  in  form  of  sharp  rugosities ; 
less  developed  asperities  on  ventral  surface  of  abdomen;  specialized 
platelets  on  venter  of  abdominal  segments  1  to  8  (poorly  developed 
on  1  to  3  and  8,  well  developed  on  4  to  7),  of  irregular  grouping  on  4 
and  5  (fig.  9,  c),  others  transverse  (fig.  9,  d).  Spiracles  each  sur- 
rounded by  narrow  irregular  area  of  membrane,  the  external  portion 
a  circular  flattened  disk  with  small  central  opening ;  trachea  tapering 
to  spiracular  opening,  with  internal  ring  (tr,  fig.  8,  &)  which  may 
function  as  closing  apparatus.  Modified  anus  (fig.  8,  e)  with  well- 
sclerotized  and  pigmented  closing  valves  (cv),  integument  at  lateral 
extremities  wrinkled,  that  posterior  and  anterior  to  valves  moderately 
pigmented,  a  short  section  of  membrane  on  internal  surface  an  appar- 
ent remnant  of  hind  gut  (hg) . 

Exuviae  of  the  first  three  instars  are  very  closely  appressed  to  each 
other  along  the  posterior  half  of  their  abdominal  sterna,  so  much  so 
that  they  can  scarcely  be  separated  without  injury  to  the  specimens. 
The  closest  attachment  appears  to  be  a  little  anterior  to  the  modified 
anal  opening  (hold-fast  organ) .  The  platelets  comprising  the  micro- 
structure  of  the  inteis:ument  on  the  lateral  and  dorsal  surfaces  of 


CRICKET    PARASITE GURNET  31 

thorax  and  abdomen,  which  in  the  fourth  instar  vary  from  tightly 
grouped  on  the  side  of  the  thorax  (fig.  9,  a)  to  narrowly  separated  on 
the  abdomen,  are  much  less  closely  grouped  in  the  third  and  second 
instars,  while  they  are  so  distant  and  reduced  in  size  in  the  first  instar 
as  to  resemble  very  widely  spaced  spiculelike  spots  of  pigment. 
Mandibles  of  the  third  and  second  instars  roughly  resemble  those 
of  the  fourth,  and  bear  sharp  teeth,  there  being  at  least  six  to  eight 
in  the  third  and  apparently  fewer  in  the  second.  The  epicranium  of 
the  first  instar  has  about  two  pairs  of  rather  large  areas  of  darker 
pigment,  but  no  mouth  parts  have  been  observed,  though  they  may 
have  escaped  detection.  There  are  10  pairs  of  spiracles  in  the  third 
and  second  instars,  with  the  pre-spiracular  tracheal  ring  poorly  defined. 
Only  six  pairs  of  spiracles  have  been  found  on  the  exuviae  of  the  first 
instar,  and  the  posterior  one  is  poorly  developed.  Each  of  the  three 
early  instars  is  brown,  darker  on  the  head.  The  venter  is  pale  in 
the  third  and  second  instars.  Approximate  lengths  of  the  exuviae  of 
the  Falls  Chuch,  Va.,  larva  are  as  follows  (in  millimeters)  :  Fourth 
6.5 ;  third  3.6 ;  second  2.3 ;  first  1.2.  Those  of  the  Marietta,  Ga.,  larva 
are  almost  identical,  but  the  Fuguay,  N.  C,  larva  is  larger,  the  three 
exuviae  measuring  as  follows :  Third  4.4 ;  second  2.6 ;  first  1.3. 

Eggs  apparently  are  broadly  elongate-oval;  fragments  are  about 
1.4  millimeters  long. 

A  basic  paper  on  the  larval  head  of  parasitic  Hymenoptera  is  that 
of  Vance  and  Smith  (1933).  Among  the  more  specialized  papers, 
those  of  G.  C.  Wheeler  (1943)  and  Beirne  (1941)  have  aided  me  most 
while  studying  the  anatomy  of  fifth-  and  fourth-instar  heads,  re- 
spectively, of  Rhopalosoma.  The  book  by  Clausen  (1940)  is  a  val- 
uable aid  toward  understanding  the  biology  and  morphology  of 
related  families,  and  it  should  be  consulted  for  references  to  the  im- 
portant work  of  J.  C.  Bridwell,  F.  A.  Fenton,  R.  C.  L.  Perkins,  F.  X. 
Williams,  F.  van  Emden,  and  others. 

Summary  and  discussion 

Among  the  biological  data  additional  to  those  brought  out  by  Hood 
(1914)  is  the  fact  that  Ha'pithus  agitator  is  an  acceptable  host  for 
Rhopalosoma  in  the  eastern  United  States.  The  effect  of  the  parasite 
on  the  internal  organs  of  the  host,  unfortunately  based  on  only  one 
dissected  cricket,  has  been  described.  Details  concerning  the  position 
of  the  parasite  egg  and  various  larval  stages  have  been  given.  Notes 
on  adult  habits,  essentially  lacking  previously,  make  it  clear  that  in 
parts  of  the  Southeast  specimens  of  Rhopalosoma  are  not  nearly  so 
scarce  as  has  been  supposed.  Like  many  another  rarity  in  collections, 
these  insects  can  doubtless  be  taken  in  numbers  by  those  familiar  with 


32  PROCEEDINGS    OF    THE    NATIONAL    MUSEUM  vol.  los 

their  appearance,  time  of  occurrence,  and  favored  habitats.  It  is  note- 
worthy that  on  a  single  day,  September  12,  1950,  Dr.  Townes  and  his 
associates  found  two  larvae,  one  on  Hapithus^  the  other  on  Orocharis. 
Specimens  belonging  to  the  University  of  Michigan  show  that  on  three 
other  occasions,  once  in  Indiana  and  twice  in  Florida,  a  collector  has 
found  two  instances  of  parasitism  in  a  single  day. 

There  remain  many  unanswered  questions  concerning  the  biology 
of  the  species  of  Rhopalosoma,  some  of  which  may  require  controlled 
cage  studies.  Stages  earlier  than  the  fourth  instar  are  still  known 
from  exuviae  and  fragments  of  eggshell,  and  first  instar  specimens 
are  especially  desirable  for  morphological  study. 

More  study  material,  including  specimens  from  northern  and  south- 
ern States,  may  show  that  more  than  one  form  of  Rho'palosoma  occurs 
in  the  United  States.  Adults  of  nearcticum  were  distinguished  from 
their  Neotropical  congeners  by  Brues  (1943)  on  rather  minor  morpho- 
logical features,  and  differences  correlated  with  geographic  distribu- 
tion might  be  better  represented  in  larvae  than  adults.  Of  the  four 
fifth-instar  specimens  examined,  the  ones  from  Georgia  and  North 
Carolina  and  the  exuviae  of  Hood's  Maryland  specimen  lack  teeth  on 
the  mandibles,  while  the  larva  from  Virginia  apparently  has  a  full 
complement  of  teeth  on  the  left  mandible  but  only  a  few  basal  ones  on 
the  right  mandible.  It  is  not  known  whether  occasional  breakage  of 
teeth  occurs  while  the  larvae  are  burrowing  in  soil  prior  to  cocoon 
formation.  Some  breakage  of  the  teeth  of  the  Virginia  larva  may 
have  resulted  from  my  manipulation  of  the  mandibles  with  needles 
before  I  realized  the  existence  of  teeth.  The  three  fifth-stage  larvae 
have  been  measured  for  maximum  head  width  and  length  of  mandibles. 
The  ratio  of  head  width  to  length  of  mandible  is  1.68, 1.86,  and  2.0  for 
the  larvae  from  Virginia,  North  Carolina,  and  Georgia,  respectively. 

Brief  examination  of  a  few  examples  of  other  families  has  sug- 
gested certain  characters  which  deserve  careful  attention  in  subse- 
quent work  of  a  comparative  nature.  A  chrysidid  larva  {Ghrysis) 
has  the  second  thoracic  spiracle  located  behind  the  groove  separating 
meso-  and  metathorax,  while  the  corresponding  spiracle  of  Rhopalo- 
soma  is  slightly  in  front  of  the  groove.  Unlike  the  condition  in 
Rhopalosoma^  the  second  spiracle  of  a  tiphiid  larva  (Oosila)  is  smaller 
than  adjacent  ones.  This  tiphiid  larva  has  two  maxillary  appendages, 
and  there  are  two  prominent  angular  projections  on  the  labium  in  the 
area  of  the  sericteries.  As  more  details  of  rhopalosomatid  life  histories 
become  available,  comparisons  with  the  situation  in  the  Tiphiidae, 
such  as  has  been  discussed  by  Clausen,  King,  and  Teranishi  (1927), 
will  perhaps  be  instructive. 

A  larva  of  one  of  the  Larridae  {Larra  analis  Fabricius,  which  at- 
tacks  mole   crickets)    has   the  head   proportionately   smaller   than 


CRICKET    PARASITE — GURNEY  33 

Rhopalosoma^  but  as  in  the  latter  the  maxilla  and  labium  have  two 
pairs  of  appendages  each.  A  conspicuous  difference  is  that  the  man- 
dible of  Larva  has  a  single  preapical  tooth  in  the  incisor  area,  and 
basally  on  a  lower  lever  there  are  two  heavy  molar  teeth,  or  "pros- 
thecae."  The  mesal  pair  of  elongate  labial  appendages  of  RhopalosoTna 
has  apparent  homalogues  in  this  larrid. 

The  mouth  parts  of  those  few  larvae  of  Bethylidae  and  Dryinidae 
that  I  have  seen  are  much  reduced  or  otherwise  specialized.  Illustra- 
tions of  dryinid  larvae  on  the  bodies  of  leafhoppers,  such  as  those 
of  Haupt  (1916)^  show  that  the  parasite  larva  sometimes  remains 
associated  with  its  exuviae  in  a  manner  suggestive  of  Rhopalosoma. 
The  genus  Harpagocryptus  Perkins,  based  on  a  species  that  Perkins 
(1908)  described  from  Queensland,  is  of  interest  in  this  connection 
because  the  larva  was  said  to  form  "a  sac  on  the  sides  of  the  abdomen 
of  small  crickets  (Trigonidiidae)."  Although  originally  referred  by 
Perkins  to  the  Dryinidae,  Bischoff  (1927,  p.  407)  called  Harpago- 
cryptus  a  bethylid,  and  Reid  (1939)  suggested  that  it  may  be  a  psam- 
mocharid.  As  with  Rhopalosoma^  a  knowledge  of  the  immature  stages 
of  Harpagocryptus  might  prove  of  great  value  when  adequate  com- 
parative studies  eventually  are  possible. 

References 

AsHMEAD,  William  H. 

1896    Rhopalosomidae,  a  new  family  of  fossorial  wasps.     Proc.  Ent.  Soc. 
Washington,  vol.  3,  pp.  303-309. 
Betkne,  Bryan  P. 

1941     A  consideration  of  the  cephalic  structures  and  spiracles  of  the  final 
instar  larvae  of  the  Ichneumonidae   (Hym.).  Trans.   Soc.  British 
Ent.,  vol.  7,  pp.  123-190,  31  figs. 
Bischoff,  H. 

1927     Biologic  der  Hymenopteren.     598  pp.,  224  figs.     Berlin. 
Blatchlet,  W.  S. 

1920  Orthoptera  of  Northeastern  America.     784  pp.,  246  figs.     Indianapolis. 
Brites,  Charles  T. 

1921  Correlation  of  taxonomic  affinities  with  food  habits  in  Hymenoptera, 

with  special  reference  to  parasitism.    Amer.  Naturalist,  vol.  55,  pp. 
134-164. 
1943     The  American  species  of  Rhopalosoma.    Ann.  Ent.  Soc.  Amer.,  vol.  36, 
pp.  310-318,  9  figs. 
Brues,  Charles  T.,  and  Melandeb,  A.  L. 

1932     Classification  of  insects.     Bull,  Mus.  Comp.  ZooL,  vol.  73,  pp.  1-672, 
1121  figs. 
Clausen,  Curtis  P. 

1940    Entomophagous  insects.     688  pp.,  257  figs. 
Clausen,  Curtis  P.,  King,  J.  L,,  and  Tebanishi,  Cho 

1927  The  parasites  of  PopilUa  japonica  in  Japan  and  Chosen  (Korea), 
and  their  introduction  into  the  United  States.  U.  S.  Dep.  Agr.  Dep. 
Bull.  1429,  1-55  pp.,  1  pi.,  35  figs. 


34  PROCEEDmGS    OF    THE    NATIONAL    MXTSEUM  vol.  io3 

CUSHMAN,  R.  A. 

1926     Some  types  of  parasitism  among  the  Ichneumonidae.     Proc.  Ent.  Soc. 
Wasliington,  vol.  28,  pp.  29-51,  51  figs. 
Haupt,  H. 

1916    Beitrage  zur  Kenntnis  der  Cicadinenfeinde.    Zeitschr.  Wiss.  Insekt- 
enb.,  vol.  12,  pp.  217-223,  fig.  13. 
Hebabd,  Morgan 

1931     The  Orthoptera  of  Kansas,     Proc.  Acad.  Nat.  Sci.  Philadelphia,  vol.  83, 
pp.  119-227,  2  figs. 

1938  Where  and  when  to  find  the  Orthoptera  of  Pennsylvania,  with  notes 

on  the  species  which  in  distribution  reach  nearest  this  state.    Ent. 
News,  vol.  49,  pp.  97-103. 
Hood,  J.  Douglas 

1914    Notes  on  the  life  history  of  Rhopalosoma  poeyi  Cresson.    Proc.  Ent. 
Soc.  Washington,  vol.  15  (1913),  pp.  145-147,  fig. 
Krombein,  K.  V. 

1951     [Rhopalosomatidae]  in  Muesebeck,  et  al.,  Hymenoptera  of  America 
north  of  Mexico.    U.  S.  Dep.  Agr.  Monogr.  2,  pp.  773-774. 
Perkins,  R.  C.  L. 

1908     Some   remarkable   Australian   Hymenoptera.     Proc.    Hawaiian   Ent. 
Soc,  vol.  2,  pp.  27-35. 
IlEHN,  J.  A.  G.,  AND  Hebabd,  Morgan 

1916     Studies  in  the  Dermaptera  and  Orthoptera  of  the  Coastal  Plain  and 
Piedmont  region  of  the  southeastern  United  States.     Proc.  Acad. 
Nat.  Sci.  Philadelphia,  vol.  68,  pp.  87-314,  3  pis. 
Reid,  J,  A. 

1939  On  the  relationship  of  the  hymenopterous  genus  Olixon  and  its  allies, 

to  the  Pompilidae  (Hym.).    Proc.  Ent.  Soc.  London,  ser.  B,  vol.  8,  pp. 
95-102,  2  figs. 
Vance,  A.  M.,  and  Smith,  H.  D. 

1933     The  larval  head  of  parasitic  Hymenoptera  and  nomenclature  of  its 
parts.    Ann.  Ent.  Soc.  Amer.,  vol.  26,  pp.  86-94,  8  figs. 
Westwood,  J,  O. 

1874    Thesaurus  entomologicus  Oxoniensis.    205  pp.,  40  pis. 
Wheeler,  George  C. 

1943    The  larvae  of  the  army  ants.     Ann.  Ent.   Soc.  Amer.,  vol.  36,  pp. 

319-332,  2  pis.,  2  figs. 
1948    The  larvae  of  the  fungus-growing  ants.     Amer.  Midi.  Nat,  vol.  40,  pp. 
664-689,  3  pis.,  3  figs. 
Wheeler,  William  Morton 

1919     The  parasitic  Aculeata,  a  study  in  evolution.     Proc.  Amer.  Philos. 
Soc,  vol.  58,  pp.  1-40. 


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Vol.   103  Washington:  1953  No.  3314 

PHOTURIS  BETHANIENSIS,  A  NEW  LAMPYRID  FIREFLY 


By  Frank  A.  McDermott 


In  August  1949,  at  Bethany  Beach  in  southern  Delaware,  some  ob- 
servations were  made  on  the  few  lampyrids  then  still  flying.  A  single 
specimen  of  Photinus  pyralis  (Linnaeus,  1767)  and  one  of  Photuris 
versicolor  (Fabricius,  1798)  were  observed,  and  a  very  few  instances 
of  a  double  greenish  flash  were  noted  over  grass  in  a  large  vacant  area 
at  the  north  end  of  the  town  and  perhaps  1,000  feet  from  the  ocean 
front.  The  insects  giving  this  flash  were  not  captured,  however,  in 
spite  of  much  pursuit,  but  another  insect  was  seen  to  fly  into  the  grass 
while  giving  the  frequently  repeated,  rather  dim  flash  characteristic  of 
females  of  Photuris.  This  one  was  taken  and  proved  to  be  a  rather 
small  female  Photuris  with  a  distribution  of  the  black  pigmentation  on 
the  pronotum  different  from  any  previously  seen.  Previous  visits 
to  Bethany  Beach  earlier  in  the  year  had  yielded  nothing  notable,  ex- 
cept the  relative  scarcity  of  lampyrids.  In  July  1951  I  again  spent  a 
few  days  at  Bethany  Beach,  and  again  saw  the  double  greenish  flash. 
This  was  considerably  later  than  the  usual  period  of  prevalence  of  P. 
versicolor.,  and  it  seemed  probable  that  this  flash  was  that  of  a  different 
species.  Several  males  were  taken,  and  one  female,  the  latter  unfor- 
tunately lost  by  accident,  but  it  may  be  noted  that  the  females  are 
apparently  relatively  scarce. 

It  differs  from  other  photurids  so  far  found  in  Delaware  by  the 
relatively  small  size,  the  distribution  of  the  pronotal  pigmentation, 
and  the  rather  distinctly  different  flash  of  the  male.  The  pronotal 
black  pigmentation,  while  rather  variable,  has  a  distinct  tendency  to 
be  enlarged  toward  the  apical  margin,  and  in  none  of  the  specimens 
did  this  spot  show  the  forms  characteristic  of  those  of  P.  versicolor., 

228423—53  35 


36  PROCEEDINGS    OF    THE    NATIONAL    MUSEUM  vol.  loa 

P.  lucricrescens  Barber,  1951,  or  P.  hebes  Barber,  1951,  all  of  which, 
had  been  collected  during  the  same  summer  in  northern  Delaware. 
The  characteristic  male  flash  is  much  like  that  of  Photinus  consangui- 
neus  LeConte,  1851,  two  short,  bright  coruscations  separated  by  a 
distinct  interval  perhaps  somewhat  longer  than  that  of  a  single  corus- 
cation, but  is  quite  greenish  as  compared  with  the  yellow  light  of 
consanguineus.  Occasionally  the  second  coruscation  may  be  much  less 
bright  than  the  first,  even  with  the  same  specimen,  and  a  few  appar- 
ently gave  only  a  single  flash  but  were  indistinguishable  from  those 
giving  the  double  flash. 

The  first  male  specimens  were  taken  on  July  11,  1951,  over  the  same 
field  where  the  single  female  was  taken  in  1949,  and  where  they  were 
still  scarce,  but  two  days  later  a  large  colony  was  found  flying  over  the 
vegetation,  largely  bayberry,  on  both  sides  of  the  road  leading  south 
from  the  town,  at  about  two  miles.  Trips  to  areas  a  mile  or  so  inland 
from  the  beach  yielded  only  Photinus  pyralis^  here  very  abundant  at 
the  time,  and  an  occasional  stray  Photuris  versicolor^  so  the  new  species 
seems  to  be  a  rather  strictly  seacoast,  almost  a  sand-dune,  form.  It 
has  not  been  seen  in  several  visits  along  the  southern  New  Jersey  coast, 
particularly  at  Stone  Harbor,  but  it  would  be  somewhat  surprising  if 
it  were  not  present  there. 

In  view  of  these  circumstances,  this  insect  is  here  described  from 
the  males  collected  in  1951,  and  the  single  female  of  1949,  as  a  new 
species. 

Photuris  bethaniensis,  new  species 

Description. — A  rather  small  lampyrid  for  a  Photuris,  9.0-10.75 
mm.  long  by  3.5-4.0  mm.  wide,  with  a  black  pronotal  mark  tending 
to  be  densest  tow^ard  the  apical  edge.  The  characteristic  male  flash  is 
two  greenish  coruscations  separated  by  a  short  interA^al. 

Head:  Width  across  eyes  2.1-2.45  mm.  in  male,  1.8  mm.  in  female; 
eye  length  1.15-1.3  mm.  in  male,  0.85  mm.  in  female.  Frons  ivory  to 
yellow,  about  0.8-1.0  mm.  wide;  interocular  margins  usually  rather 
divergent.  Mandibles  fairly  large,  0.65-0.8  mm.  across  in  closed  posi- 
tion; proportionately  larger  in  the  female.  Maxillary  palpi  dark 
brown,  apical  joint  conoidal ;  labial  palpi  lighter  brown,  apical  joint 
mitten-shaped.     Clypeus  dark  brown,  more  or  less  tridentate. 

Antennae:  Dark  brown,  5.0-6.25  mm.  long  in  male,  4.25  mm.  in 
female ;  joints  may  all  have  pale  bases,  or  this  may  be  limited  to  basal 
three  joints  in  the  female. 

Pronottjm:  Scutate,  sometimes  extended  apically,  2.0-2.5  by 
2.55-3.0  mm. ;  rounded  posterior  angles  may  or  may  not  be  appreciably 
produced ;  red  pigmentation  may  extend  to  the  basal  edge ;  black  spot 
of  variable  shape,  and  characteristically  mainly  toward  the  apex,  or 


NEW    LAMPYEID    FIREFLY — McDERMOTT  37 

at  least  the  larger  portion  of  it  apical,  the  longitudinal  portion  being 
usually  merely  an  irregular  and  sometimes  interrupted  line,  and  but 
little  developed  along  the  basal  margin. 

SCUTELLUM  AND  MESONOTAL  PLATES  I    White  Or  yelloW. 

Elytra  :  7.0-8.0  by  1.6-2.0  mm. ;  ground  color  brown  to  nearly 
black;  on  each  a  white  to  yellow  vitta,  oblique,  usually  extending  to 
about  three-quarters  the  elytral  length ;  wide  yellowish  lateral  borders 
which  give  a  slightly  oval  contour;  narrow  sutural  borders. 

Ventral  thoeacic  surface  :  Brown. 

Tekglfes  :  Tergites  1  to  6  brown ;  tergite  7  lighter  or  only  partially 
inf  uscate ;  tergite  8  pale  to  white. 

Sternites  :  2  to  5  usually  brown,  sometimes  becoming  darker  in  this 
order;  sternite  5  may  be  yellow;  sternites  6  and  7  luminous  in  the 
male ;  and  sternite  8  with  a  median  posterior  point,  which  may  be  very 
short.  The  luminous  area  in  the  female  differs  from  the  pattern  seen 
in  versicolor^  being  a  transverse  bilobed  area  medially  very  narrow,  on 
segment  6,  and  a  relatively  large  elliptical  area  on  segment  7. 

Legs  :  Long ;  tibial  spurs  long,  pattern  0-2-2. 

Aedeagus  :  In  the  one  dissected,  2.25  mm.  long,  identical  in  appear- 
ance with  that  of  P.  lucicrescens  and  the  other  photurids  described  by 
Barber.^ 

Type  aifid  paratypes. — USNM  61469. 

Type  locality. — Bethany  Beach,  Del. 

Material  examined. — Holotype  (male)  and  five  paratypes,  collected 
by  McDermott  at  type  locality,  July  11, 1951;  paratype  (female),  col- 
lected by  McDermott  at  type  locality,  August  1949. 

Remarks. — Photuris  hethaniensis  is  probably  a  seacoast  form;  it 
has  not  been  noted  elsewhere.  The  flash  suggests  P.  fairchildi  Barber, 
1951,  but  the  latter  is  a  larger  species  with  narrow  elytral  borders. 


*  Barber,  H.  S.     North  American  fireflies  of  the  genus  Photuris.     Smithsonian  Misc.  Coll., 
vol.  117,  No.  1,  pp.  1-58,  3  figs.,  1951. 


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Vol.    103  Washington:  1953  No.  3315 

DISTRIBUTION,  GENERAL  BIONOMICS,  AND  RECOGNITION 
CHARACTERS  OF  TWO  COCKROACHES  RECENTLY  ESTAB- 
LISHED  IN  THE  UNITED  STATES 


By  Ashley  B.  Gurney^ 


Introduction 


In  contrast  to  the  large  numbers  of  cockroach  species  that  occur 
in  a  great  variety  of  natural  habitats  in  most  tropical  countries,  only 
about  55  are  now  recorded  from  the  entire  United  States.  With  the 
exception  of  those  occurring  in  a  few  localities,  mainly  in  the  South- 
western States,  our  roaches  are  well  known  to  entomologists.  In  the 
northern  third  of  the  country  there  are  few  native  roaches  living  in 
fields  or  woodlands,  and  most  of  the  better-known  species  are  estab- 
lished adventives  which  are  rather  closely  associated  with  human 
habitations,  having  been  brought  there  by  artificial  means.  The  recent 
establishment  in  the  Northeastern  States  of  two  potentially  important 
roaches  new  to  this  country  is  therefore  of  decided  interest. 

Beginning  in  1948  and  continuing  each  subsequent  year,  specimens 
of  Ectohius  livens  (Turton),  the  spotted  Mediterranean  roach,^  have 
been  collected  at  Falmouth,  Mass.,  under  outdoor  conditions,  coming 
to  houses  at  night  and  also  occurring  on  fresh  vegetables.  Because 
of  these  habits  and  the  wide  natural  occurrence  of  this  roach  in  Europe 
and  adjacent  areas,  it  seems  quite  probable  that  a  slow  spread  in  the 
United  States  may  take  place,  and  that  from  time  to  time  questions 
regarding  its  identity  and  importance  will  arise. 

1  Bureau  of  Entomology  and  Plant  Quarantine,  Agricultural  Research  Administration, 
U.  S.  Department  of  Agriculture. 

2  No  generally  accepted  common  name  seems  to  be  applied  to  E.  livens  in  Europe,  and 
this  name  is  considered  suitable  for  American  use  in  view  of  the  species'  appearance  and 
the  region  of  its  dominant  Old  World  occurrence. 

39 

228354— 53— —1 


40  PROCEEDINGS    OF    THE    NATIONAL    MUSEUM  vol.  io3 

A  second  species  not  previously  known  to  breed  in  the  United  States 
is  LeiLCophaea  maderae  (Fabricius),  the  Madeira  roach.  Unlike  the 
spotted  Mediterranean  roach,  L.  maderae  is  well  known  because  it  is 
frequently  transported  with  articles  of  commerce  in  tropical  countries. 
It  has  many  times  been  intercepted  by  quarantine  inspectors  at 
United  States  ports  and  has  occasionally  been  carried  to  inland  cities, 
where  it  failed  to  survive.  In  the  autum  of  1950  it  was  found  to 
be  established  in  the  basements  of  New  York  City  buildings  occupied 
by  people  who  had  come  from  Puerto  Rico.  Although  its  native 
home  is  evidently  West  Africa,  it  apparently  was  introduced  to  the 
West  Indies  more  than  100  years  ago.  The  Madeira  roach  could 
probably  live  outdoors  in  tropical  Florida  and  perhaps  at  the  other 
southern  extremities  of  the  United  States,  but  it  is  more  likely  to  be 
important  as  a  domestic  pest  in  the  well-heated  buildings  of  our  larger 
cities.  One  has  only  to  recall  the  increasing  importance  of  the  brown- 
banded  roach  Supella  supellectiliuTii  (Serville)  in  this  country  during 
comparatively  recent  years  to  realize  the  potential  importance  of 
Leucophaea  maderae.  Apparently  a  native  of  Africa  that  has  been 
spread  by  commerce  to  most  of  the  warmer  parts  of  the  world,  S. 
supellectilium  was  first  reported  from  the  United  States  in  1903,  based 
on  specimens  from  Key  West,  Fla.  It  was  not  noted  as  a  pest  here 
until  found  in  a  private  home  in  Nebraska  in  1929,  but  since  then 
has  rapidly  gained  recognition  as  one  of  our  principal  pest  roaches 
(see  Back,  1937,  and  Gould  and  Deay,  1940) . 

This  paper  is  intended  to  aid  in  the  identification  of  these  two 
roaches  and  to  supply  such  available  information  on  distribution  and 
habits  as  is  likely  to  be  helpful  to  subsequent  studies  of  these  species 
in  the  United  States.  A  third  species,  Nauphoeta  cinerea  (Olivier), 
was  found  to  be  established  in  Florida  after  this  manuscript  was 
prepared.     It  is  briefly  noted  on  page  46. 

Genus  Ectohius  Stephens,  1835 
Ectobius  livens  (Turton),  1800?  (spotted  Mediterranean  roach) 

Figure  10,  a-d:  Plate  2,  Figures  3-6 

Blatta  livida  Fabricius,  Entomologia  systematicae,  vol.  2,  p.  10,  1793.  (Pre- 
occupied by  Blatta  livida  De  Geer,  Memoirs  pour  servir  a  I'histoire  des  In- 
sectes,  vol.  3,  p.  538,  pi.  44,  fig.  6, 1773.) 

Blatta  livens  Turton.  A  general  system  of  nature  .  .  .  ,  vol.  2,  p.  529,  1800? 
(New  name  for  Blatta  livida  Fabricius.) 

EctoMus  lividus  Fabricius,  Stephens,  Illustrations  of  British  entomology,  Mandi- 
bulata,  vol.  6,  p.  48, 1835. 

Ectohius  livens  (Turton) ,  Ramme,  Mitt.  Zool.  Mus.  Berlin,  vol.  27,  pp.  34-45,  pi.  10, 
fig.  3, 1951. 

Nomenclatural  coTnments. — ^Until  1951  Ectobius  lividus  was  usually 
applied  to  this  species.    The  name  E.  livens  has  been  resurrected  by 


COCKROACHES    NEW    TO    UNITED    STATES — GURNEY  41 

Kamme  (1951,  p.  34)  who  says  that  Dr.  K.  Princis  of  Lund,  Sweden, 
informed  him  that  livida  Fabricius  was  preoccupied  and  that  Turton 
proposed  livens  on  page  526,  vol.  2,  of  his  1806  edition  of  Linne's  Sys- 
tema  Naturae.  Turton  published  two  editions  of  an  English  transla- 
tion of  the  thirteenth  (Gmelin)  edition  of  this  work.  These  editions 
are  listed  in  the  1903-1915  Catalogue  of  the  Books,  Manuscripts,  Maps 
and  Drawings  in  the  British  Museum  (Natural  History)  (vol.  3, 1910, 
p.  1128).  I  have  verified  their  dates  and  contents  by  an  examination 
of  the  entire  7-volume  1806  edition  in  the  Library  of  Congress  and  of 
volume  2  of  the  4-volume  first  edition  in  the  library  of  the  U.  S.  Na- 
tional Museum.  The  first  Turton  edition  appeared  from  1800  to  1802. 
Volume  2,  containing  the  genus  Blatta^  is  dated  1800  and  is  reasonably 
sure  to  have  appeared  no  later  than  1802.  The  second  Turton  edition 
appeared  in  1806,  the  first  4  volumes  being  re-issues  of  the  former 
edition  with  slight  changes  of  the  title  pages.  On  the  page  indicated 
by  Ramme  (p.  526)  Forftcula  livida^  an  earwig,  is  treated. 

Prior  to  Ramme's  monograph,  the  identities  of  the  British  species  of 
Ectohius  were  confused.  Thus  the  species  treated  by  Lucas  (1920,  p. 
78)  as  pers-picillaris  Herbst  is  in  reality  livens.  This  has  been  clarified 
by  Blair  (1934,  1935).  Princis  (1936)  has  explained  that  a  Lapland 
record  of  lividus  (i.  e.,  livens)  actually  was  based  on  material  of 
lapponicus  (Limiaeus).  Hebard  (1943,  p.  12)  states  that  Australian 
records  of  lividus  are  incorrect.  Taxonomists  now  differentiate  the 
species  of  Ectobius  on  rather  minute  characters,  especially  on  the  basis 
of  the  dorsal  abdominal  gland  and  genitalia  of  males.  Ramme  ( 1949 ) 
has  described  E.  siculus  from  Sicily,  differing  from  livens  in  having 
black  legs  and  antennae.  E.  finoti  Chopard  of  Algeria  (see  Chopard, 
1943,  p.  21)  is  much  like  livens,  having  the  small  tegminal  spots  between 
the  veins  and  with  the  dorsal  gland  bearing  a  rounded  tubercle.  The 
genus  EctoMus  (genotype:  lappoQiicns)  in  a  modern  sense  is  much 
more  restricted  than  formerly.  In  the  older  literature,  species  of 
EctoMus  were  reported  from  many  parts  of  the  world.  Certain  of 
these  are  now  known  to  belong  to  other  genera.  For  instance,  many 
years  ago  several  Nearctic  roaches  were  referred  to  Ectobius.  Rehn 
(1931,  pp.  306-374)  and  Hebard  (1943)  have  discussed  the  generic 
limits  of  Ectobius:  Rehn  (p.  308)  has  explained  the  nature  of  the 
toothed  tarsal  claws ;  the  teeth  were  evidently  overlooked  by  Hebard 
(p.  12). 

Recognition  features. — The  most  important  features  enabling  rec- 
ognition of  Ectobius  livens  are:  (1)  Small  size,  over-all  length  about 
8  to  9  millimeters ;  (2)  small,  dark  spots  on  the  veins  of  the  front  wing 
(tegmen) ;  (3)  conspicuous  intercalated  triangle  at  apex  of  wing  (itr, 
fig.  10,  b) ',  (4)  claws  of  unequal  length,  the  posterior  claw  of  each 
pair  much  longer  than  anterior  one;  (5)  in  males  the  characteristic 
shape  of  the  dorsal  abdominal  glandular  depression  and  the  sub- 


42  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  los 

genital  plate  (fig.  10,  «,  c)  ;  (6)  conspicuous  spines  on  ventroposterior 
margin  of  each  femur. 

Of  the  foregoing  features,  the  second  and  fifth  are  specific  for  the 
separation  of  this  roach  from  other  species  occurring  in  the  United 
States  and  the  others  are  highly  confirmatory  when  in  combination. 

The  native  Nearctic  species  most  likely  to  be  mistaken  ^  for  E.  livens 
are  Chorisoneura  texensis  Saussure  and  Zehntner,  of  Texas  and  the 
Southeast,  and  members  of  the  genus  Cariblatta.  Both  have  claws  of 
equal  length,  and  Cariblatta  lacks  a  well-developed,  intercalated  tri- 
angle. C.  texensis  has  a  large  intercalated  triangle,  but  has  no  con- 
spicuous spines  along  the  posterior  margins  of  the  femora.  The  best- 
known  outdoor  roaches  of  the  Eastern  States  belong  to  the  genus  Par- 
coblatta;  all  are  larger  than  livens  and  lack  an  intercalated  triangle. 

Identification  of  the  Massachusetts  specimens  of  livens  was  aided 
by  comparisons  made  at  the  Academy  of  Natural  Sciences  of  Phila- 
delphia with  authentic  French  and  Swiss  material,  given  to  the  Acad- 
emy by  the  Swiss  entomologist  Henri  de  Saussure. 

Description. — Adults  :  General  form  as  in  plate  2,  figure  6 ;  tegmina 
and  wings  covering  abdomen ;  width  between  eyes  on  vertex  about  1^ 
times  length  of  first  antennal  segment  (female),  slightly  less  in  male; 
all  femora  with  conspicuous  spines  on  ventroposterior  margins ;  ven- 
troposterior margin  of  front  femur  with  three  or  four  well-spaced, 
strong  spines  along  part  slightly  basad  of  middle,  the  more  apical  one 
often  smaller  than  others,  followed  by  10  to  15  small,  delicate  spines 
(some  of  which  are  frequently  broken)  extending  in  a  regular  row 
nearly  to  apex,  a  long  curved  apical  spine  closely  preceded  by  a  straight 
subapical  one  about  three- fourths  as  long ;  tarsal  segments  one  to  four 
each  with  a  tiny,  round  pulvillus  (membranous  pad)  ;  segment  5  with 
large,  broad  apical  arolium  between  conspicuously  unequal  tarsal 
claws,  anterior  claw  about  two-thirds  length  of  posterior  one,  longer 
claw  with  about  four  spinelike  inner  teeth  along  basal  half,  teeth  on 
smaller  claw  poorly  developed.  Tegmen  with  costal  veins  simple, 
cubital  veins  (discoidal  sectors)  strongly  oblique,  apex  rather  sharply 
rounded.    Wing  with  distinctive  intercalated  triangle  (itr,  fig.  10, 

General  color  pale  yellowish;  tegmina  and  marginal  area  of  pro- 
notum  transparent,  marked  with  reddish  brown  spots,  on  the  tegmen 
these  spots  located  on  the  veins ;  eyes  dark  brown  to  black ;  face  yellow 
to  reddish  orange ;  vertex  and  upper  part  of  face  usually  with  several 
spots  of  darker  orange;  coxae  and  femora  unspotted;  tibiae  with 
dark  brown  at  bases  of  some  spines;  venter  of  abdomen  with  poorly 

*The  standard  taxonomie  reference  on  Nearctic  Blattarla  Is  that  of  Hebard  (1917).  A 
recent  key  to  genera  is  by  John  W.  H.  Rehn  (1950).  The  European  species  of  EctoMus 
were  reviewed  by  Ramme  (1923). 


COCKROACHES    NEW    TO    UNITED    STATES — GURNEY  43 

developed  longitudinal  rows  of  dark  brown  siibmarginal  and  median 
spots. 

Measurements  (in  millimeters)  of  representative  specimens:  Over- 
all length,  including  folded  tegmina,  female  9,  male  8.5;  pronotum, 
female  2.1,  male  1.9 ;  tegmen,  female  7,  male  7.2.  Width  of  pronotum, 
female  3,  male  2.9.     No  significant  size  variation  has  been  noted. 

Nymphs  :  Nymphs  (pi.  2,  figs.  3,  4)  ranging  in  body  length  from 
2.5  to  4.5  millimeters  have  been  examined.  They  are  yellow,  with 
conspicuous  dark  reddish  brown  spots  on  the  dorsal  surface  and  on  the 
vertex  and  upper  part  of  the  face.  Dark  submarginal  longitudinal 
streaks  appear  on  the  thorax.  The  venter  of  the  abdomen  is  dark 
brown  submarginally  and  medially. 

Oothecae:  Two  oothecae  have  been  examined  (pi.  2,  fig.  5).  They 
are  dark  brown  and  measure  2.3  millimeters  in  length.  The  convex 
margin  is  weakly  crenulate  and  there  are  faint  transverse  indications 
of  the  eggs  contained. 

Distribution. — Three  American  localities  are  known  for  Ectobius 
livens,  all  in  Massachusetts:  Falmouth,  Manomet,  and  Plymouth. 
Falmouth,  the  first  site  discovered  here,  is  situated  near  Woods  Hole 
and  the  Cape  Cod  Canal,  on  the  southern  side  of  the  base  of  Cape  Cod. 
In  October  1951,  after  this  manuscript  had  been  prepared,  I  ex- 
amined 3  males  and  4  females  of  livens  collected  at  Manomet,  Plym- 
outh County,  Mass.,  by  Mrs.  S.  P.  Graeff  and  submitted  by  Dr.  Ells- 
worth H.  Wheeler  of  the  University  of  Massachusetts.  Manomet  is 
on  the  coast  of  Cape  Cod,  about  20  miles  north  of  Falmouth.  The 
specimens  were  collected  before  August  13,  1951,  apparently  just 
prior  to  that  date.  Mrs.  Graeff  wrote,  "Our  summer  cottage  is  pretty 
well  overrun  with  them.  Painters  told  me  they  were  even  on  the  roof. 
The  cottage  is  in  the  woods  and  the  underbrush  is  close.  I  think  this 
is  the  third  summer  we  have  had  them.  I  suspect  they  like  damp 
spots,  they  scurry  under  shingles." 

An  adult  of  livens  was  submitted  July  28,  1952,  by  a  Plymouth 
housewife  who  reported  that  her  house  and  shrubbery  were  becoming 
infested. 

This  roach  is  widely  distributed  in  the  southern  and  central  portions 
of  western  Europe,  occurring  in  Germany,  the  Netherlands,  Belgium, 
France,  Switzerland,  Italy,  Portugal,  Spain,  and  southern  England. 
It  has  also  been  reported  as  far  eastward  in  the  Mediterranean  area 
as  Asia  Minor,  but  the  records  I  have  seen  were  published  prior  to 
the  modern  work  dating  from  Ramme's  1923  monograph  of  Ectobius 
and  are  not  dependable.  Chopard  (1943,  p.  21)  has  reported  it  from 
Algeria  and  Tunisia.  I  have  found  no  records  of  its  occurrence  in 
the  Azores.  Fabricius  originally  described  the  species  from  France, 
and,  in  the  absence  of  type  material,  Eamme  in  1923  designated  a  male 


44  PROCEEDINGS    OF    THE    NATIONAL    MUSEUIVI  vol.  io3 

neotype  from  Montpellier  in  southern  France.  Two  varieties  of 
livens  have  been  recognized,  chopardi  Adelung  described  from  France 
in  1916,  and  minor  Kamme  described  from  the  Island  of  Elba  in  1923. 
These  varieties  are  based  primarily  on  the  amount  of  dark  color  on 
the  lower  surface  of  the  abdomen,  minor  being  quite  dark  except  near 
the  lateral  margins,  only  the  submarginal  dark  spots  being  well  de- 
veloped in  chopardi^  and  the  submarginal  spots  being  poorly  or  moder- 
ately developed  in  typical  livens.  The  species  of  Ectohius  are  well 
known  for  color  variation,  the  color  of  tegmina  and  abdomen  often 
being  recessive  (pale)  or  intensive  (dark).  In  view  of  this  situation, 
it  remains  to  be  proved  whether  chopardi  and  minor  are  valid  sub- 
species or  if  they  are  forms  without  real  significance.  The  center  of 
distribution  for  the  genus  Ectohius^  as  outlined  by  Rehn  (1931,  pp. 
314—318),  was  apparently  the  Mediterranean  Basin,  with  a  secondary 
center  developed  in  Africa  south  of  the  Sahara  Desert. 

Biology. — The  original  Massachusetts  specimens  of  Ecto'hius  livens 
examined  consist  of  2  males,  16  females,  9  nymphs,  and  2  oothecae. 
The  males  were  taken  June  21,  1948,  and  the  other  specimens  in  late 
August  and  September  1950.^  It  may  be  significant  that  males  were 
found  only  in  the  spring  and  early  summer.  Lucas  (1928)  reported 
that  in  England  males  of  E.  lapponicus  disappeared  about  midsum- 
mer, the  females  persisting  longer  than  males,  and  nymphs  appearing 
in  late  summer  and  fall  preparatory  to  hibernation.  No  detailed 
life  history  studies  of  livens  in  Europe  have  come  to  my  attention. 
Brown  (1952)  has  reported  observations  on  the  related  E.  panzeri 
Stephens  in  England,  which  occurs  there  mainly  on  sand  dunes  near 
the  coast.  Adults  die  in  the  fall,  the  males  disappearing  first,  but 
nymphs  overwinter  and  do  not  reach  full  maturity  until  early  August 
of  the  following  year. 

The  Falmouth  collections  were  made  on  a  farm  located  beside  a 
small  salt  river  about  a  mile  from  the  shore.  Mr.  Flint  found  speci- 
mens in  a  large  variety  of  situations,  specimens  occurring  most  con- 
sistently beneath  loose  lichens  on  oak  trees,  and  crawling  on  trees  and 
houses  at  night — the  latter  in  the  vicinity  of  lights.  Some  were 
found  under  baskets,  in  buckets,  or  on  Swiss  chard  and  other  vege- 
tables, or  were  taken  by  general  sweeping  (see  Flint,  1951). 

In  England  E.  livens  occurs  only  in  the  extreme  southern  counties — 
there  found  on  trees,  among  bracken  ferns,  under  dead  leaves,  and 
flying  actively  in  hot  sunshine  (Lucas,  1920,  p.  80;  Burr,  1936,  p.  44). 
Chopard  ( 1947,  p.  37)  says  livens  is  very  common  in  the  French  woods, 
and  he  gives  a  colored  illustration  (pi.  2,  fig.  28) . 

■*  AU  of  this  material  was  collected  by  Oliver  S.  Flint,  Jr.,  a  student  at  the  University 
of  Massachusetts.  I  am  much  indebted  to  Mr.  Flint  for  the  enthusiasm,  with  which  he 
sought  additional  material  once  the  character  of  his  Initial  captures  was  pointed  out  to 
him,  as  well  as  for  his  diligence  in  seeking  clues  to  the  origin  of  the  introduction. 


COCKROACHES    NEW   TO    UNITED    STATES — GURNEY  45 

Source  of  introduction. — The  Massachusetts  colony  of  Ectohius 
livens  is  probably  the  result  of  an  accidental  introduction  from  western 
Europe  or  the  Mediterranean  area.  A  good  many  people  of  Portu- 
guese ancestry  live  in  Falmouth  and  the  vicinity,  and  visits  to  Portu- 
gal and  the  Azores  are  frequent.  Seeds  and  occasional  shipments 
of  fruit  are  brought  back,  but  quarantine  regulations  prevent  wide- 
spread movements  of  plant  products.  Since  livens  occurs  in  Portugal, 
as  reported  by  Seabra  (1942,  p.  19),  the  possibility  of  an  introduc- 
tion from  that  part  of  the  Mediterranean  seems  the  most  likely. 

Probable  importance. — Ectohius  livens  is  not  likely  to  develop  into 
a  major  pest  or  to  live  in  buildings  except  occasionally.  As  a  nuis- 
ance pest  in  the  vicinity  of  gardens  and  dwellings,  it  may  be  expected 
to  attract  attention  at  intervals  and  to  be  submitted  to  entomologists 
for  identification  and  advice,  as  is  now  true  of  male  specimens  of 
Parcohlatta  which  are  attracted  to  lights  and  which  occasionally  re- 
main in  houses  for  short  periods. 

So  far  as  known,  species  of  Ectohius  are  not  economically  impor- 
tant in  Europe.  E.  lapponicus  has  often  been  stated  to  be  a  pest  of 
dried  fish  in  Swedish  Lappland,  but  Gaunitz  (1935,  1936)  found 
that  laffonicus  does  not  occur  indoors  and  probably  feeds  on 
vegetable  matter. 

Genus  Leucophaea  Brunner,  1865 
Leucophaea  maderae  (Fabricius),  1793  (Madeira  roach) 

Figure  10,  e-h;  Plate  2,  Figxjbes  1,  2 

Blatta  maderae  Fabricius,  Entomologia  systematicae,  vol.  2,  p.  6, 1793. 
Leucophaea  maderae  (Fabricius),  Rehn,  Trans.  Amer.  Ent.  Soc,  vol.  29,  p.  283, 
1908. 

Nomenclatural  comments. — The  name  Rhyparohia  maderae  is  cur- 
rently used  by  certain  workers,  especially  Europeans,  as  exemplified 
by  Chopard  (1943,  p.  45)  in  his  monograph  of  North  African  Orthop- 
tera.  Leucophaea  was  proposed  by  Brunner  (1865,  pp.  272,  278) 
as  a  subgenus  of  Panchlora.^  with  four  included  species.  One  of  the 
four,  maderae^  was  designated  type  of  Leucophaea  by  Rehn  (1903, 
p.  282).  Rhyparohia  was  proposed  by  Krauss  (1892,  p.  165),  mono- 
typic  for  maderae.  Thus  Rhyparohia  is  an  isogenotypic  synonym  of 
Leucophaea.  The  use  of  Rhyparohia  was  doubtless  encouraged  by  the 
action  of  Kirby  (1904,  pp.  150-151)  who,  evidently  unaware  of  Rehn's 
1903  designation,  designated  Blatta  surinamensis  Linnaeus  type  of 
Leucophaea.  As  explained  by  Hebard  (1917,  p.  309),  Kirby  also  ap- 
parently objected  to  the  name  Pycnoscelus  Scudder,  1862,  to  which 
surinamensis  belongs,  because  it  was  originally  based  on  an  immature 
specimen.    Kirby's  latter  view  is  not  supported  by  the  rules  of  nomen- 


46  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  yol.  los 

clature  but,  regardless  of  that  aspect  of  the  matter,  the  validity  of 
Leucophaea^  with  maderae  its  type,  is  clear. 

Recognition  features. — The  most  important  features  in  the  recog- 
nition of  Leucophaea  maderae  are :  (1)  Large  size,  overall  length  about 
38  to  53  millimeters;  (2)  pronotal  shape,  and  general  color  markings 
as  illustrated  (pi.  2,  figs.  1,  2) ;  (3)  no  strong  spines  along  posterior 
margins  of  femora  except  one  short  apical  spine  on  each  hind  and 
middle  femur;  (4)  in  males  the  characteristic  shape  of  the  specialized 
organ  on  the  dorsal  surface  of  the  second  abdominal  segment  and  of 
the  subgenital  plate  (fig.  10,  e,  h.). 

Relatively  few  Nearctic  roaches  are  as  large  as  L,  maderae.,  and  none 
of  them  is  likely  to  be  confused  with  it.  Blaherus  craniifer  Burmeister, 
illustrated  by  Hebard  (1917,  pi.  8,  fig.  6),  is  the  largest  roach  in  the 
United  States,  being  much  larger  than  maderae.  The  species  of 
Perijylaneta  are  quite  differently  marked,  and  have  numerous  heavy 
spines  on  the  posterior  margins  of  the  femora.  Both  Eurycotis 
floridana  (Walker),  of  the  Southeast,  and  Hemihldbera  tenehricosa 
Rehn  and  Hebard,^  of  the  Florida  Keys,  have  short,  lobate  tegmina 
and  wings  which  are  vestigial  or  lacking. 

A  relative  of  L.  maderae  recently  established  in  Florida  is 
Nauphoeta  cinerea  (Olivier).  It  superficially  resembles  maderae.^ 
though  it  is  smaller,  its  average  length  being  about  23  millimeters. 
The  femora  are  proportionately  stouter  and  the  tegmina  scarcely  ex- 
tend beyond  the  apex  of  the  abdomen,  while  the  posterior  margin  of 
the  pronotum  is  transversely  subtruncate,  a  median  projection  being 
scarcely  evident.  Zimmerman  (1948,  p.  94)  and  Rehn  (1945,  p.  274) 
have  given  photographs  of  cinerea.  This  species  is  apparently  a  na- 
tive of  East  Africa  now  widely  distributed  in  the  Tropics  by  com- 
merce and  often  intercepted  at  United  States  ports.  Its  potential 
importance  is  suggested  by  Illingworth  (1942),  who  described  its  oc- 
currence in  alarming  numbers  about  the  feed  room  of  the  poultry 
plant  at  the  University  of  Hawaii.  He  also  mentioned  its  occurrence 
in  heated  buildings  in  England  and  Germany,  though  no  records  of 
permanent  establishment  in  those  countries  have  come  to  my  attention. 

In  early  December  1951  material  of  cinerea  was  received  from  Jo- 
seph Gross  of  Tampa,  Fla.,  who  stated  that  it  was  widely  and  thor- 
oughly established  about  Tampa,  especially  in  feed  mills.  Later,  in 
August  1952,  Mr.  Gross  contributed  a  series  of  34  adults  and  32  nymphs 

s  IlemiMaiera  tenebricosa  was  described  from  the  West  Indies,  including  Nassau  in  the 
Bahamas,  by  Rehn  and  Hebard  (1927,  p.  271).  Caudell  (1931)  recorded  an  adult  pair 
collected  at  Key  Largo.  Fla.,  by  E.  A.  Popenoe  in  1896.  Caudell's  unpublished  notes  dis- 
close subsequent  discussions  of  this  record  with  Mr.  Popenoe's  son,  who  explained  that  his 
father  was  collecting  insects  primarily  for  Kansas  State  College  during  the  1896  trip  and 
that  in  view  of  conditions  at  Key  Largo  at  that  time  this  roach  was  probably  an  established 
species  rather  than  an  intercepted  adventive.  A  number  of  roaches  are  now  known  to 
be  native  to  the  northern  Bahamas  and  extreme  tropical  Florida,  so  the  occurrence  of 
tenehricosa  in  the  Florida  Keys  Is  not  illogical. 


COCKROACHES    NEW    TO    UNITED    STATES — GURNEY 


47 


Figure  10. — Structural  details  of  Ectobius  livens  (Turton)  and  Leucophaea  maderae  (Fab- 
ricius).  a-d,  E.  livens:  a.  Dorsal  view  of  apical  half  of  cleared  male  abdomen;  Z»,  wing; 
c,  ventral  view  of  subgenital  plate  and  associated  cerci,  male;  d,  apical  portion  of  genital 
hook,  male,  e-h,  L.  maderae:  e,  Dorsal  view  of  second  abdominal  segment,  male,  cleared 
and  in  alcohol;/,  dorsal  view  of  apical  half  of  cleared  male  abdomen;  g,  genital  hook  of 
male,  enlarged;  h,  ventral  view  of  subgenital  plate  and  associated  structures,  male,  dry 
preparation.  Drawn  from  specimens  as  follows:  a-d,  from  Massachusetts  specimen;  e, 
from  Puerto  Rico  specimen;  /  and  g,  from  specimen  collected  in  a  Tampa,  Fla.,  warehouse; 
A,  from  New  York  City  specimen.  (A-9,  apodemes  of  sternum  9;  ca,  oblique  carinae  of 
glandular  specialization;  gh,  genital  hook;  gld,  glandular  depression;  itr,  intercalated 
triangle;  me,  mesal  extension;  st,  stylus;  sap,  supra-anal  plate;  sgp,  subgenital  plate.) 
228354—53 2 


48  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  io3 

to  the  U.  S.  National  Museum.  I  inquired  especially  as  to  the  apparent 
permanence  of  the  infestation,  and  on  June  24, 1952,  a  colleague  of  Mr. 
Gross',  W.  B.  Gresham,  Jr.,  replied,  "It  seems  evident  to  us  that 
Nauphoeta  is  in  Florida  to  stay.  They  appear  to  be  well  adapted  and 
quite  numerous  in  the  locations  where  we  have  noted  them.  All  in- 
festations noted  have  been  within  commercial  buildings  engaged  in  the 
manufacture  of  animal  feeds.  I  myself  have  not  noted  any  out  of 
doors."  Notes  on  the  Florida  infestations  by  Ratcliffe  (1952)  and 
Gresham  (1952)  have  appeared. 

Description. — Adults  :  General  form  as  in  plate  2,  figure  1 ;  tegmina 
and  wings  extending  beyond  apex  of  abdomen  about  one-fifth  their 
length ;  width  between  eyes  on  vertex  about  two-thirds  length  of  first 
antennal  segment ;  a  shallow,  scarcely  wrinkled  transverse  depression 
in  interocular  area  where  eyes  are  closest ;  ventroposterior  margin  of 
middle  and  hind  femur  each  with  short,  apical  spine;  conspicuous 
spines  on  posterior  femoral  margins  otherwise  lacking;  ventropos- 
terior margin  of  front  femur  with  continuous  row  of  delicate,  seta- 
like spines,  these  slightly  shorter  toward  apex  of  femur  and  a  few 
noticeably  longer  ones  near  base ;  front  tibia  with  two  elongate  groups 
of  setae  along  margins;  pulvillus  on  each  tarsal  segment,  elongate 
on  segments  1  and  5;  claws  equal,  unarmed  ventrally,  with  sparse 
setae  dorsally ;  pronotum  obtusely  angulate  laterally,  posterior  margin 
with  rounded  median  production;  tegmen  with  cubital  veins  (dis- 
coidal  sectors)  oblique;  wing  (see  J.  W.  H.  Eehn,  1951,  pi.  9,  fig.  102) 
with  well-developed  axillary  and  cubital  fields,  intercalated  triangle 
lacking.  Dorsal  surface  of  abdomen  simple  in  female,  specialized  in 
male;  second  tergum  of  male  (fig.  10,  e)  with  median  glandular  de- 
velopment consisting  of  paired  oblique  carinae  (ca)  and  a  brief  longi- 
tudinal carina  centered  on  a  trifid  papilla  which  largely  covers  a 
small  glandular  depression  (gld)  ;  supra-anal  plate  transverse,  with 
median  emargination  of  posterior  margin,  general  shape  more  quadrate 
when  dry  (fig.  10,  h)  than  when  preserved  in  alcohol  (fig.  10,  /)  ;  sub- 
genital  plate  unspecialized  in  female,  slightly  asymmetrical  in  male, 
with  recurved  lateral  hooks  near  styli ;  cleared  male  abdomen  with 
apodemes  of  sternum  9  (subgenital  plate)  (A9)  united  anteriorly, 
genital  hook  (gh)  and  additional  sclerotized  elongate  rod  associated 
with  an  irregular  genital  mass;  genital  hook  (fig.  10,  g)  enclosed  in  a 
membranous  sheath. 

General  color  pale  brown,  the  tegmen  and  pronotum  marked  with 
dark  brown  as  illustrated  (pi.  2,  fig.  1)  ;  remainder  of  body  and  ap- 
pendages largely  dark  brown ;  clypeus  and  most  of  labrum  pale ;  ab- 
dominal sterna  with  oblique  submarginal  marks  of  brownish-black; 
terga  with  more  heavily  pigmented  areas  as  shown  (fig.  10,  /).  Base 
of  genital  hook  with  pigmented  transverse  lines,  the  apical  portion 
much  darker. 


U.  S.  NATIONAL  MUSEUM 


PROCEEDINGS.  VOL.  103  PLATE  2 


1,  2,  Leucophaea  maderae  (Fabricius);  1,  Adult  female,  collected  in  Manila,  Philippine 
Islands  (length,  47  mm.):  2,  last  nymphal  stage,  intercepted  at  Washington,  D.C. 

3-6,  Ectohius  livens  (Turton),  all  from  Falmouth,  Massachusetts:  3,  4,  Nymphal  stages; 
5,  cotheca;  6,  adult  female  (length,  9  mm.). 


COCKROACHES    NEW    TO    UNITED    STATES — GURNEY  49 

Measurements  (in  millimeters)  :  Overall  body  length,  including 
folded  tegmina,  female  45  to  53,  male  38  to  45 ;  pronotum,  female  9  to 
10.3,  male  7.5  to  8.5 ;  tegmen,  female  37  to  46,  male  32  to  38.5.  Width 
of  pronotum,  female  13.3  to  16,  male  11.5  to  14. 

Nymphs:  Nymphs  (pi.  2,  fig.  2)  ranging  from  7.5  to  32  millimeters 
in  body  length  have  been  examined.  The  dorsal  surface  of  the  thorax 
and  abdomen  is  sharply  rugose,  with  short,  microscopic  spines,  these 
especially  conspicuous  along  the  posterior  margins  of  segments; 
much  smaller  and  less  evident  spines  on  ventral  surface  of  abdomen. 
General  color  reddish  brown ;  each  tergum  with  a  submarginal  dark 
spot  near  the  base,  the  basal  half  of  the  exposed  lateral  margin  black- 
ish and  the  apical  third  yellowish. 

Distnhution. — In  the  United  States  Leucophaea  maderae  is  known 
to  be  established  only  in  the  Harlem  section  of  New  York  City.*'  Many 
times  each  year  inspections  made  at  shipping  centers  detect  the  species 
with  plant  products  and  other  articles  coming  both  from  the  American 
Tropics  and  the  Old  World.  The  distribution  of  maderae  in  warm 
countries  is  very  wide,  but  somewhat  irregular,  depending  upon  the 
vagaries  of  commerce.  The  specific  and  the  common  names  are  de- 
rived from  the  fact  that  this  roach  was  first  described  from  Madeira, 
a  small  island  now  belonging  to  Portugal  and  located  in  the  Atlantic 
Ocean  about  400  miles  west  of  Morocco.  Eehn  ( 1937,  pp.  56-58 ;  1945, 
p.  273)  has  reasoned  (by  a  detailed  analysis  of  the  present  occurrence 
of  Tnaderae  and  its  congeners)  that  West  Africa  is  the  native  home, 
from  which  commerce  (in  early  centuries — that  of  the  slave  trade  in 
particular)  has  carried  it  to  Madeira,  the  West  Indies,  Brazil,  and 
elsewhere.  Ecuador,  Colombia,  Venezuela,  the  Guianas,  Brazil,  and 
Argentina  are  known  to  have  established  colonies  of  maderae^  as  well 
as  Panama,  Costa  Rica,  and  most  of  the  West  Indian  islands.  Old 
World  distribution  outside  of  Africa  is  more  spotty  and,  to  my  knowl- 
edge, the  species  is  not  yet  recorded  from  India  or  Australia — even 
though  it  is  known  from  Java,  the  Philippines,  Hawaii,  and  Fiji. 
Doubtless,  7naderae  now  breeds  in  manj^^  coastal  areas  from  which 
there  are  no  records  and  it  will  continue  to  spread  as  conditions  permit. 

Biology. — In  New  York  City  Leucophaea  maderae  has  been  "found 
in  some  abundance  in  the  basement  of  an  apartment  of  the  Harlem 
section,"  and  an  observer  also  writes  ^  "From  their  abundance  and  the 
presence  of  all  stages  of  the  insect  ....  I  would  conclude  that  they 


•Davis  (1940)  reported  at  a  meeting  held  December  15,  1938,  that  a  specimen  of 
L.  maderae  taicen  in  the  Bird  House  of  Bronx  Park,  New  York  City,  was  the  first  record 
of  tliis  species  in  New  York  State.  In  the  absence  of  further  information,  there  is  a  strong 
likeliliood  that  the  specimen  in  question  was  an  escaped  adventive. 

'  Specimens  from  the  described  infestation  were  submitted  by  Ralph  E.  Heal.  Technical 
Director  of  the  National  Pest  Control  Association.  Grateful  acknowledgment  is  made  of 
Dr.  Heal's  cooperation  in  obtaining  information  on  the  Harlem  Infestation. 


50  PROCEEDINGS    OF    THE    NATIONAL    MUSEUM  vol.  io3 

are  breeding  in  this  house."     It  was  also  noted  that  the  Harlem  roaches 
appeared  rather  sluggish  in  their  movements. 

In  the  absence  of  biological  studies  of  L.  maderae  in  this  country  it 
is  instructive  to  review  the  observations  of  life  history  and  habits 
that  have  been  made  elsewhere,  particularly  the  papers  of  Illingworth 
(1915),  Sein  (1923),  and  Pessoa  and  Correa  (1928). 

As  is  true  of  species  of  PeHplaneta^  growth  in  the  Madeira  roach 
is  slow.  The  first  molt  may  occur  1  to  4  months  after  birth.  Pinned 
nymphs  reared  by  Illingworth  in  Hawaii  and  now  deposited  in  the 
U.  S.  National  Museum  suggest  that  there  are  six  nymphal  stages, 
though  the  number  is  likely  to  vary,  and  in  cultures  it  may  sometimes 
be  difficult  to  observe  because  exuviae  reportedly  are  eaten  immedi- 
ately after  each  molt.  Pessoa  and  Correa  report  only  four  "meta- 
morphoses," but  that  number  of  molts  is  almost  certainly  too  small 
for  a  normal  growth  cycle.  The  body  length  in  millimeters  of  speci- 
mens of  various  ages  collected  by  Illingworth  is  as  follows:  1  day, 
7.5;  1  month,  7.5;  4  months,  16;  5  months,  18.5;  6  months,  24.5;  7 
months,  29 ;  8  months,  30.7 ;  9  months,  32.  Food  and  environmental 
conditions  are  doubtless  important  influences  on  growth.  Sein  notes 
maturity  being  attained  in  220  days,  but  that  a  year  is  often  required, 
while  Pessoa  and  Correa  indicate  that  adults  are  obtained  in  a  year, 
with  16  to  18  months  the  longest  observed  time.  In  Sein's  cultures 
the  first  nymphs  appeared  4  months  after  their  parents  matured. 
Inmiediately  following  birth,  nymphs  usually  hide  beneath  the 
mother  during  the  day.  According  to  Pessoa  and  Correa,  this  makes 
the  adult  restless  and  active  in  contrast  to  its  usual  slow  gait. 

In  a  general  paper  emphasizing  the  adaptability  and  value  of 
Leucophaea  Trmderae  as  an  experimental  animal,  Scharrer  (1951) 
comments  that  it  thrives  on  a  diet  of  apples,  carrots,  and  dogfood. 
She  reports  that  there  are  30  to  35  young  every  3  months,  that  there 
is  an  average  of  8  molts,  and  that  life  expectancy  is  up  to  2V^  years. 
Dr.  Scharrer  has  observed  that  tumors  which  resemble  malignant 
cancer  of  higher  animals  develop  in  various  parts  of  the  roach  body 
following  the  removal  of  the  corpora  allata  and  the  corpora  cardiaca. 
That  such  an  injury  to  the  nervous  system  has  produced  tumors  has 
very  interesting  and  practical  implications  for  further  research. 
The  name  "woodroach"  that  Dr.  Scharrer  applied  to  maderae  is  a 
questionable  choice,  since  the  name  is  most  often  used  for  native 
Nearctic  species  of  Parcohlatta. 

Leucophaea  is  one  of  several  roach  genera  which  are  viviparous; 
that  is,  the  eggs  are  enclosed  in  a  delicate  membranous  sac  which  nor- 
mally ruptures  before  extrusion  from  the  mother's  body,  or  immedi- 
ately thereafter,  so  that,  in  effect,  the  young  are  born  alive.  This  is 
in  contrast  to  the  habit  of  most  roaches  which  form  a  heavily-sclero- 
tized,  dark-colored  capsule  or  ootheca,  usually  of  characteristic  shape, 


COCKROACHES    NEW    TO    UNITED    STATES — GURNEY  51 

which  may  be  carried  protruding  from  the  body  for  some  time  prior 
to  hatching,  or  be  deposited  loosely  or  surrounded  with  a  matrix. 

The  viviparity  exhibited  generally  takes  a  special  form  known  as 
ovoviviparity.  This  occurs  in  several  well-known  genera,  especially 
Panchlora^  Pycnoscelus^  Nauphoeta^  and  Leucophaea^  and  Chopard 
(1938,  p.  218)  notes  six  subfamilies  of  roaches  (according  to  the  long- 
used  classification)  in  which  viviparity  occurs.  J.  W.  H.  Rehn  (1951)' 
has  recently  proposed  a  new  classification,  based  on  wings,  and  it  is 
significant  that  many  of  the  viviparous  genera  fall  in  what  he  terms 
the  epilamproid  complex.  In  addition  to  Chopard,  Shelf ord  (1907), 
Karny  (1924,  pp.  3-10),  and  Rau  (1941)  have  discussed  viviparity 
among  roaches.  Hagan  (1941;  1951)  has  described  the  female  re- 
productive system  of  the  viviparous  Diploptera^  and  Chopard  (1950), 
that  of  Gromphadorhina. 

lUingworth  and  Sein  both  recognized  that  Leucophaea  maderae  is 
viviparous,  but  it  appears  that  Pessoa  and  Correa  observed  unusual 
or  abnormal  instances  of  the  Qgg  sac  being  deposited,  as  they  wrote  of 
a  capsule  being  placed  in  the  darkest  corner  of  a  rearing  box  and  the 
first  young  appearing  20  days  later.  In  the  following  paragraphs  I 
have  attempted  to  explain  this  apparent  lack  of  agreement. 

Several  preserved  adult  females  of  L.  maderae  received  for  identifi- 
cation during  recent  years  have  exhibited  an  elongate  sac  about  20 
millimeters  long  protruding  from  the  end  of  the  abdomen.  In  cer- 
tain cases  the  eggs  (varying  up  to  about  40  in  number),  directed 
transversely  with  respect  to  the  mother's  body  and  arranged  in  two 
rows,  were  undeveloped,  but  in  others  nymphs  ready  for  hatching  were 
visible — in  fact,  rupturing  of  the  sac  had  sometimes  occurred.  Rehn 
(1937,  p.  62)  has  described  his  experience  in  the  Belgian  Congo  with 
the  related  L.  grandis  (Saussure) .  A  female  confined  in  a  bottle  gave 
birth  to  20  living  young.  The  number  of  young  of  tnaderae  produced 
at  one  time,  according  to  published  records,  ranges  from  25  to  32. 

The  related  Pycnoscelus  surinamensis  (Linnaeus)  is  known  to  be 
viviparous,  but  the  literature  concerning  its  habits  suggests  a  lack  of 
uniformity  in  birth  or  hatching.  Watson  (1929,  p.  58)  commented  on 
an  egg  capsule  being  deposited  but  the  eggs  failing  to  hatch.  Caudell 
(1925)  also  maintained  cultures  of  P.  surinamensis,  and  mentioned 
young  born  alive  and  oothecae  seldom  if  ever  protruding  from  the 
female  abdomen.  In  a  somewhat  fuller  account,  Zappe  (1918)  gave 
the  opinion  that  young  of  surinamensis  are  either  born  alive  or  hatch 
from  eggs  within  24  hours.  Eggs  were  often  laid  in  soil,  but  were  not 
observed  to  hatch.  When  females  were  injured  or  excited,  they  often 
deposited  poorly  developed  egg  masses  that  did  not  hatch.  Similarly, 
lUingworth  (1942)  reported  that  the  eggs  of  Nauphoeta  cinerea  were 
usually  kept  in  the  body  until  living  young  appeared,  but  that  confined 
individuals  sometimes  produced  aborted  egg  masses  that  were  ex- 


52  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.103 

truded  but  which  dried  up  without  hatching  when  the  delicate  enclos- 
ing membrane  was  exposed  to  the  air.  These  experiences  suggest  that 
some  of  the  protruding  eggs  of  L.  niaderae  that  I  have  noted  are  the 
result  of  handling  when  the  specimens  were  collected  and  that  occa- 
sional Qgg  masses  may  be  deposited  under  certain  conditions  by  this 
ordinarily  viviparous  species. 

In  his  important  recent  paper,  Chopard  (1950)  has  discussed  the 
anatomy  and  development  of  Gromfhadorhina  laevigata  Saussure 
and  Zehntner,  of  Madagascar.  He  observed,  in  the  course  of  rearings 
conducted  in  Paris,  that  the  female  of  this  viviparous  species  extrudes 
the  soft  ootheca  nearly  to  its  full  extent,  then  it  is  drawn  into  an 
incubating  pouch  where  the  eggs  undergo  incubation  for  approxi- 
mately 70  days  prior  to  the  appearance  of  the  young.  Occasionally 
the  ootheca  is  completely  extruded,  with  the  result  that  desiccation 
occurs  and  no  hatching  takes  place.  I  am  indebted  to  my  colleague 
R.  E.  Snodgrassfor  pointing  out  the  significance  of  Chopard's  obser- 
vations. They  explain  how  the  ootheca  is  transferred  from  the  uterus 
to  the  incubating  chamber  in  the  case  of  Gromphadorhina.  Possibly 
the  same  habit  occurs  in  certain  other  viviparous  genera,  and  some 
protruding  oothecae  that  have  been  seen  may  represent  a  stage  in  this 
normal  act.  In  discussing  the  transfer  of  the  developing  eggs  in 
the  viviparous  Difloptera^  Hagan  (1951,  p.  299)  states  that  the 
oocytes  pass  from  the  ovarioles  to  the  lower  end  of  the  common  ovi- 
duct, where  "they  are  directed  by  the  ovipositor  from  the  genital 
chamber  ventrally  into  the  open  end  of  the  uterus." 

In  Brazil,  mating  of  Leucofhaea  maderae  occurs  mainly  during  the 
warm  and  rainy  season,  according  to  Pessoa  and  Correa,  who  say  that 
copulation  may  occupy  20  to  30  minutes  and  takes  place  with  the  pair 
end  to  end  facing  in  opposite  directions.  Prior  to  mating,  the  female 
is  described  as  opening  her  wings  and  drawing  them  along  the  ground, 
at  the  same  time  producing  a  sound  by  vibrating  them.  The  scent 
gland  on  the  dorsum  of  the  male  presumably  is  attractive  to  the  female 
at  mating  time,  but  its  function  is  not  definitely  known. 

lUingworth  stated  that  when  disturbed  the  Madeira  roach  stridu- 
lates  very  noticeably,  and  he  believed  the  sound  is  produced  by  rubbing 
the  posterior  margin  of  the  pronotum  over  the  mesonotum.  Accord- 
ing to  Chopard  (1938,  p.  286),  several  roaches  stridulate  delicately  by 
rubbing  the  border  of  the  pronotum  upon  the  mesonotum  or  upon  the 
strongly  denticulate  base  of  the  costal  vein  of  the  tegmen.  To  test  the 
possibilities  of  stridulation,  I  have  relaxed  dry  specimens  of  both  sexes 
of  L.  maderae^  then  manipulated  the  body  parts  with  my  fingers.  A 
low,  squeaking  sound  is  consistently  obtained  in  either  sex  by  rubbing 
the  lateroposterior  margin  of  the  pronotum  on  the  basal  costal  margin 
of  the  tegmen.     Both  surfaces  are  heavily  sclerotized,  turned  to  oppose 


COCKROACHES    NEW    TO    UNITED    STATES — GURNEY  53 

each  other,  nonpubescent,  and  are  finely  rugose  so  as  to  make  stridula- 
tion  possible. 

In  tropical  regions  where  it  is  established,  Z.  maderae  is  definitely  a 
domiciliary  species,  though,  like  certain  other  roaches  often  associated 
with  man,  it  is  capable  of  living  apart  from  him  in  a  purely  wild  state. 
From  available  records,  however,  it  is  not  clear  how  frequently  it  has 
been  collected  entirely  unassociated  with  man-made  surroundings. 
According  to  Alfken  (1904,  p.  565)  more  than  50  years  ago  a  German 
collector,  H.  Schauinsland,  in  1896  or  1897  found  mfiderae  in  native 
huts  on  the  Hawaiian  Island  of  Molokai,  commerce  already  having 
carried  it  to  certain  Pacific  areas.  However,  in  spite  of  this  long  estab- 
lishment in  the  Hawaiian  group,  no  mention  of  its  occurrence  in  fields 
is  made  by  Williams  ( 1931 ) .  In  July  1950,  in  the  Venezuelan  State  of 
Aragua,  Dr.  Ernst  Schwarz  ^  collected  it  coming  to  lights  in  fair  num- 
bers, but  whether  it  was  breeding  in  the  adjacent  forest  or  associated 
with  nearby  habitations  is  not  known.  Likewise,  specimens  collected 
by  the  late  C.  F.  Baker  in  the  Philippines  before  1927  bear  the  collect- 
ing label  "Mt.  Maquiling,"  and  they  may  have  bred  under  entirely  nat- 
ural surroundings.  However,  as  I  saw  during  my  visit  to  Mt.  Maquil- 
ing in  1945,  at  the  base  and  on  the  lower  slopes  there  are  ample  oppor- 
tunities for  this  roach  to  remain  associated  with  man.  The  species 
frequently  flies  actively  and  has  often  been  taken  at  lights,  including 
those  on  porches,  in  field  camps,  or  of  automobiles.  On  the  other  hand, 
Sein  states  that  Leucophaea  maderae  does  not  fly  much  when  indoors 
at  night  as  part  of  an  infestation.  L.  puerilis  Rehn,  considered  its 
closest  relative,  is  a  strictly  endemic,  forest  type  in  West  Africa. 

Source  of  introduction. — The  New  York  City  infestation  is  reported 
to  be  localized  in  apartment  buildings  occupied  by  people  from  Puerto 
Rico  and,  since  there  has  been  a  good  deal  of  movement — much  of  it 
by  air — from  that  island  in  the  past  few  years,  there  is  little  doubt  that 
Puerto  Rico  is  the  source  of  the  infestation. 

Probable  importance. — In  Puerto  Rico  the  Madeira  roach  most  often 
occurs  in  fruit  stores  and  markets.  It  is  especially  fond  of  grapes.  It 
is  considered  very  gregarious  and  develops  large,  localized  colonies. 
Wolcott  (1950,  p.  43)  records  about  a  bushel  having  been  swept  from 
one  store.  Warehouses  and  other  buildings  are  often  infested.  When 
handled  or  otherwise  disturbed,  Leucophaea  maderae  produces  an  odor 
described  as  especially  offensive.  Houses  infested  by  the  species  usu- 
ally do  not  contain  Periplaneta  or  Blattella.  There  is  a  strong  possi- 
bility that  maderae.,  if  allowed  to  spread,  will  gradually  develop  into  a 
serious  pest  in  our  larger  cities.  In  the  light  of  its  wide  occurrence  in 
the  Bahamas  and  other  West  Indian  islands,  tropical  Florida  would 
probably  be  a  suitable  habitat,  either  outdoors  or  in  buildings  without 
central  heating. 

« Associated  with  the  Venezuela  Plague  Mission  (Commander  J.  M.  Amberson,  D.S.N., 
Dr.  Ernst  Schwarz,  and  Mrs.  Schwarz). 


54  PROCEEDESTGS    OF    THE    NATIONAL   MUSEUM  vol.108 

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1940.  The  biology  of  six  species  of  cockroaches  which  inhabit  buildings. 

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Hagan,  Haeold  R. 

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2  figs. 
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1917.  The   Blattidae  of  North  America  north  of  the  Mexican  boundary. 

Mem.  Amer.  Eut.  Soc,  No.  2,  pp.  1-284,  10  pis.,  1  fig. 
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1892,  Systematische  Verzeichnis  der  Canarischen  Dermapteren  und  Orthop- 
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56  PROCEEDENGS    OF    THE    NATIONAL   MUSEUM  vol.  loa 

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U.  S.  GOVERNMENT  PRfNTINS   OFFICE:  1993 


PROCEEDINGS   OF   THE   UNITED   STATES   NATIONAL   MUSEUM 


SMITHSONIAN  INSTITUTION 
U.  S.  NATIONAL  MUSEUM 


Vol.   103  Washington  :  1953  No.  3316 

BITING  MIDGES  OF  THE  HELEID  GENUS  STILOBEZZIA  IN 

NORTH  AMERICA 


By  Willis  W.  Wirth  ' 


Siilobezzia  Kieffer,  1911,  is  a  rather  large,  widespread  genus  of 
predaceous  midges  which  has  reached  its  greatest  development  in 
the  warmer  parts  of  the  New  World.  For  North  America  11  species 
are  listed  in  Johannsen's  recent  list  (1943b);  this  total  is  now  raised 
to  18  by  the  present  new  records  and  species,  and  no  doubt  many 
more  await  discovery.  The  species  of  Stilobezzia  are  rather  easily 
characterized,  and  all  the  Nearctic  species  known  to  Johannsen  were 
correctly  placed  by  him,  with  the  exception  of  S.  uncinata  Johannsen, 
1943,  which  I  believe  to  represent  the  female  of  Parabezzia  petiolata 
Malloch,  1915. 

The  types  of  the  species  described  here  are  in  the  U.  S.  National 
Museum,  the  collections  of  which  furnished  the  bulk  of  the  material 
studied.  I  am  indebted  to  Dr.  Henry  K.  Townes,  of  North  Carolina 
State  College,  for  the  loan  of  specimens  from  his  collection,  and  to 
Mrs.  Elisabeth  C.  Beck  and  J.  A.  Mulrennan,  of  the  Florida  State 
Department  of  Health,  for  their  kindness  in  sending  a  large  series  of 
light  trap  collections  from  which  I  sorted  most  of  the  Florida  specimens. 

Family  Heleidae 

Genus  Stilobezzia  Kieflfer,  1911 

Diagnosis. — Body  rather  slender  and  nearly  bare.  Eyes  bare. 
Female  antenna  with  segments  3-10  oval,  11-15  long  and  cylindrical; 


'  Bureau  of  Entomology  and  Plant  Quarantine,  Agricultural  Research  Administration,  U.  S.  Depart' 
ment  of  Agriculture. 

232991—53 1  57 


58  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  103 

male  antenna  with  well-developed  plumes.  Mesonotum  usually  with 
few  or  no  long  bristles;  humeral  pits  present  but  usually  inconspicuous. 
All  legs  slender;  femora  without  spines  or  (in  Eukraiohelea)  with  a 
few  on  forelegs;  basitarsus  and  fifth  segment  with  or  without  ventral 
spines;  fourth  segment  cordiform,  fifth  not  enlarged;  female  claws 
large  and  very  unequal  or  a  single  long  claw  with  basal  barb  on  each 
leg,  male  claws  small  and  equal;  empodium  absent.  Wing  rather 
long;  microtrichiae  present;  macro trichiae  present  toward  tip  (in 
Neostilobezzia)  or  absent;  costa  extending  to  two-thirds  wing  length 
or  more;  usually  two  radial  cells,  second  much  larger  than  first, 
which  may  be  entirely  absent  (in  Eukraiohelea)  or  greatly  reduced; 
intercalary  fork  distinct;  crossvein  r-m  more  or  less  oblique;  median 
fork  with  long  petiole,  M2  not  interrupted  at  base;  anal  vein  straight, 
not  interrupted  in  middle.  Male  genitalia  with  basistyles  simple  or 
with  lobe  on  inner  margins ;  dististyles  long ;  ninth  sternite  very  short ; 
ninth  tergite  usually  rounded  caudad  with  setigerous  apicolateral 
lobes;  aedeagus  reduced  to  a  pair  of  oblique  lateral  sclerites;  parameres 
a  pair  of  large,  highly  sclerotized,  greatly  modified,  submedian  proc- 
esses with  flaring  basal  apodemes. 

Keys  to  the  Nearctic  species  of  Stilobezzia 

Females 

1.  Only  one  long  radial  cell  present;  fore  femur  with  2  or  3  ventral  spines 

(subgenus  Eukraiohelea) 1.  elegantula    (Johannsen) 

Two  radial  cells  present ;  fore  femur  unarmed 2 

2(1).  Wing  with  macrotrichiae  at  tip  (subgenus  Neostilobezzia) 3 

Wing  entirely  bare  (subgenus  Stilobezzia) 5 

3  (2).  Body  hairs  very  strong  and  erect;  wings  hyaline;  legs,  scutellum,  and 

halteres  yellow 3.  lutea  (Malloch) 

Body  hairs  soft  and  decumbent;  midlegs  and  hind  legs  brown 4 

4(3).  Wing  infuscated  brown;  scutellum  and  halteres  yellowish. 

4.  stonci,  new  species 
Wing  hyaline;  scutellum  and  halteres  brownish 2.  fuscula  Wirth 

5  (2).  Wing  with  2  or  3  dark  spots  on  anterior  margin 6 

Wing  hyaline,  unmarked 9 

6  (5).  Shining  yellow;  mesonotum  polished  yellow,  with  pair  of  longitudinal 

brown  bands,  anterior  spine  absent;  legs  yellow,  narrow  apices  of  hind 
femur  and  tibia  brown;  wing  hyaline,  with  brown  spots  at  crossvein 

and  apex  of  second  radial  cell 9.  coquilletti  Kieffer 

.  Mesonotum  bro^yn  with  silvery,  pruinose  areas  and  prominent  anterior 
spine;  legs  extensively  brown  banded  or  punctate;  wing  brownish, 
anterior  margin  with  2  or  3  dark  spots 7 

7  (6).  Femora  and  tibiae  extensively  yellow,  with  small  brown  spots  at  bases  of 

hairs;  wing  hyaline,  with  two  brown  spots;  abdomen  mostly  yellowish 

above 14.  punctipes,  new  species 

Femora  and  tibiae  brown  with  yellowish  rings;  wing  infuscated,  with  2 
or  3  brown  spots  and  a  subapical  pale  area;  abdomen  black  above  with 
narrow,  distal,  silvery  pruinose,  segmental  bands 8 


HELEID    MIDGES,    GENUS    STILOBEZZIA — WIRTH  59 

8  (7) .  Three  dark  wing  spots  including  one  at  apex  of  first  radial  cell;  fifth  tarsal 

segment  with  ventral  batonnets 15.  rabelloi  Lane 

Two  dark  wing  spots,  none  at  apex  of  first  radial  cell;  fifth  tarsal  segment 
without  ventral  batonnets 6.  beckae,  new  species 

9  (5) .  Mesonotum  entirely  polished  black 10 

Mesonotum  yellow,  green,  or  brown,  not  entirely  black 13 

10  (9).  Abdomen  pohshed  black 11 

Abdomen  green 12 

11  (10).  Midfemora  and  hind  femora  black  except  at  extreme  bases. 

5.  antennalis  (Coquillett) 
Legs  uniformly  yellow 7.  bicolor  Lane 

12  (10).  Fifth  tarsal  segment  with  ventral  batonnets.  _  10.  diversa  (Coquillett) 

Fifth  tarsal  segment  without  ventral  batonnets. 

12.  pallidiventris  (Malloch) 

13  (9).  Shining  green;  third  or  fourth  abdominal  tergites  extensively  black.     14 

Yellow  or  brown,  not  green;  abdomen  without  black  bands  or  spots.     15 

14  (13).  Mesonotum  uniformly  pale  green;  distal  fourth  of  hind  femur  black. 

18.  viridis  (Coquillett) 

Mesonotum  largely  dark  brown;  hind  femur  with  only  spot  at  apex 

black 11.  glauca  Macfie 

15  (13).  Mesonotum  shining  brown;  wing  with  a  beadlike  swelling  near  apex  of 

second  radial  cell 16.  sybleae,  new  species 

Mesonotum  pruinose  brown;  wing  venation  normal 16 

16  (15).  Mesonotum  dark  brown,  with  pattern  of  large  pruinose  gray  spots; 

scutellum  dark;  tibiae  with  sub-basal  pale  rings..  13.  pruinosa  Wirth 
Mesonotum  uniform  light  grayish  brown;  tibiae  unbanded 17 

17  (16).  Scutellum  and  femora  yellow 8.  bulla  Thomsen 

Scutellum  and  femora  brown 17.  thomscnae,  new  species 

Male  genitalia 

1.  Lateral  sclerites  of  aedeagus  greatly  reduced  or  absent,  instead  a  trans- 
verse sclerite  between  inner  margins  of  basistyles  beyond  base 2 

Lateral  sclerites  of  aedeagus  well  developed  and  oblique,  with  bases  at 
inner  ventral  margins  of  basistyles 3 

2  (1).  A  large  rounded  lobe  at  distal  third  of  inner  margins  of  basistyles; 

transverse  sclerite  with  pointed,  curved  ends;  apicolateral  lobes  of 

ninth  tergite  widely  spaced  and  triangular 8.    bulla  Thomsen 

Basistyles  simple;  transverse  sclerite  with  blunt  ends;  apicolateral  lobes 
fingerlike  and  close  together 17.  thomsenac,    new   species 

3  (1).  Lateral  sclerites  of  aedeagus  sinuate,  irregularly  bent,  or  apex  forked; 

basistyle  without  lobe,  or  if  one  present  it  bears  a  strong  spine;  apico- 
lateral lobes  of  ninth  tergite  usually  slender  and  close  together 4 

Lateral  sclerites  of  aedeagus  straight,  at  least  on  distal  half;  basistyle 
with  mesal  lobe;  apicolateral  lobes  usually  low  and  rounded 10 

4  (3).  Basistyle  with  a  strong  spine  midway  on  inner  margin,  arising  from  a  low 

lobe  also  bearing  a  fine  hair;  aedeagus  with  strong  subapical  tooth 
on  outer  side,  appearing  forked  at  end;  parameres  long  and  slender 

with  long  pointed  tips 2.  fuscula  Wirth 

Basistyle  simple ;  aedeagal  sclerites  without  strong  tooth 5 


60  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  103 

5  (4).  Parameres   greatly   expanded   from    bases,    with    obliquely    truncated 

apices  bearing  a  short  lateral  tooth  and  a  long,  curved,  distal  tooth. 

4.  stonei,  new  species 
Parameres  broadest  at  bases  or  without  truncated  apices 6 

6  (5).  Stems  of  parameres  very  slender  and  rodlike 7 

Stems  of  parameres  bladelike 9 

7  (6).  Apices   of  parameres    slightly    expanded,   palplike;    aedeagal   sclerites 

sinuate,  stouter  toward  apices;  ninth  sternite  with  deep,  quadrate, 
mesal  notch ;  dististyles  very  broad,  with  truncate  apices. 

1.  elegantula    (Johannsen) 

Apices  of  parameres  sharp-pointed  and  bent  ventrad;  aedeagal  sclerites 

bent  more  than  twice 8 

8  (7).  Membrane  of  ninth  sternite  bare;  tips  of  aedeagal  sclerites  recurved 

hooklike;  dististyles  very  slender 6.  beckae  new  species 

Membrane  spiculate;  tips  of  aedeagel  sclerites  snoutlike  and  crossing 
each  other;  dististyles  stout 14.  punctipes,  new  species 

9  (6).  Parameres  with  an  extra  lateral  pair  of  straight,  bladelike  arms  half 

as  long  as  inner  arms ;  membrane  of  ninth  sternite  bare. 

15.  rabelloi  Lane 

Parameres  simple,  without  lateral  arms,  expanded  subapically,  with  tips 

abruptly  narrowed;  membrane  spiculate 3.  lutea  (Malloch) 

10  (3).  Parameres  with  knobbed  apices  over  twice  as  broad  as  the  very  slender, 

stalklike  stems;  basistyles  very  large  and  globular;  dististyle  very 

slender  to  base 18.  viridis  (Coquillett) 

Parameres  otherwise,  without  knobbed  apices 11 

11  (10).  Parameres  with  greatly  swollen,  conical  bases  and  very  slender,  tapered 

apices 12 

Parameres    with    stems    of   subequal    diameter    or    swollen    towards 
apices 13 

12  (11).  Parameres  with  basal  0.6  swollen,  the  pointed  apices  abruptly  bent 

towards  base  on  outer  side 10.  diversa  (Coquillett) 

Parameres  with  only  basal  0.2  swollen,  distal  portions  consisting  of  very 
slender,  curved,  saber-shaped  blades  crossing  each  other  in  middle. 

16.  sybleae,  new  species 

13  (11).  Parameres  nearly  straight,   with  apices  abruptly  narrowed  and  bent 

over 14 

Parameres  sinuate  or  with  broad,  toothlike,  lateral,  subapical  expan- 
sions      15 

14  (13).  Parameres  with  apices  bent  outward 7.  bicolor  Lane 

Parameres  with  apices  bent  inward 11.  glauca  Macfie 

15  (13).  Lobe  of  basistyle  quadrate,  as  long  as  broad;  parameres  slender  and 

markedly  sinuate;  dististyles  greatly  curved. 

5.  antennalis  (Coquillett) 

Lobe  of  basistyle  consisting  of  only  a  hump  on  inner  margin ;  parameres 

very  broad  and  stout ;  dististyle  nearly  straight 16 

16  (15).  Each  paramere  with  broad,  toothlike,  lateral,  subapical  expansion  about 

half  as  broad  as  length  of  dististyle,  with  a  slender,  ventrally  curved 
spine  from  apex  on  inner  margin;  dististyle  tapered. 

9.  coquilletti  Kieflfer 

Each  paramere  swollen  slightly  midway  to  apex  without  subapical  lateral 

expansion;  dististyle  with  broad,  blunt  apex 13.  pruinosa  Wirth 


HELEID    MIDGES,    GENUS    STILOBEZZIA — WIRTH  61 

Table  1. — Proportions  of  segments  of  hind  legs  of  female  Stllobezzia  species. 


Species 


Cx 

Tr 

F 

Ti 

Ti 

T2 

Ta 

T, 

20 

10 

60 

60 

30 

10 

5 

4 

25 

10 

60 

65 

32 

15 

6 

4 

40 

15 

90 

90 

40 

20 

9 

6 

25 

10 

70 

70 

30 

18 

7 

5 

20 

12 

55 

55 

28 

12 

3 

3 

30 

15 

75 

75 

40 

18 

6 

5 

15 

7 

40 

35 

17 

10 

4 

3 

15 

7 

35 

35 

15 

8 

3 

2 

30 

15 

80 

85 

40 

18 

5 

5 

13 

6 

35 

32 

16 

6 

3 

3 

20 

10 

65 

70 

33 

15 

4 

4 

20 

10 

50 

50 

28 

10 

4 

4 

30 

15 

80 

80 

40 

20 

6 

4 

40 

15 

85 

95 

45 

25 

10 

5 

15 

7 

45 

50 

25 

12 

3 

2 

20 

10 

55 

55 

28 

12 

3 

3 

9. 
10. 
11. 
12. 
13. 
14. 
15. 
16. 
17. 


elegant  ula 

fuscula 

lutea 

stonei 

antennalis 

beckae 

bicolor 

bulla 

coquilletti 

diversa 

glauca 

pallidiventrio 

pruinosa 

punctipes 

rabelloi 

sybleae 

viridis 


12 

8 

15 

10 


10 
5 

13 
5 

10 


10 

15 

5 

8 


Subgenus  Eukraiohelea  Ingram  and  Macfie,  1921 

Eukraiohelea  Ingram  and  Macfie,  Ann.  Trop.  Med.  Parasit.,  vol.  15,  p.  347,  1921. — 
Johannsen,  Arch.  IlydrobioL,  Suppl.,  vol.  9,  p.  430,  1931;  Journ.  New  York 
Ent.  Soc,  vol.  42,  p.  344,  1934.— Macfie,  Ann.  Trop.  Med.  Parasit.,  vol.  34, 
p.  22,  1940.— Tokunaga,  Tenthredo,  vol.  3,  p.  344,  1940.— Johannsen,  Ann. 
Ent.  Soc.  Amer.,  vol.  36,  p.  781,  1943;  Bull.  Bishop  Mus.  No.  189,  p.  190,  1946. 
(Genotype,  Eukraiohelea  africana  Ingram  and  Macfie,  1921;  designated  by 
Macfie,  1940.) 

Diagnosis. — First  radial  cell  absent,  the  r-m  crossvein  nearly  form- 
ing a  straight  line  with  Ri,  the  latter  arising  just  a  little  more  toward 
base  of  wing;  fore  femora  with  2  or  3  ventral  spines  but  not  swollen; 
male  aedeagus  with  hyaline  posterior  membrane,  parameres  long  and 
very  slender. 

Discussion.' — Johannsen  (1934)  placed  Eukraiohelea  as  a  subgenus 
of  Parabezzia  Malloch,  1915.  However,  as  shown  by  a  comparison 
of  the  male  genitalia  of  P.  petiolata  Malloch,  1915,  with  those  of  the 
four  species  of  Eukraiohelea  whose  males  have  been  described,  the 
latter  are  much  more  closely  related  to  Stilobezzia.  The  following 
species  have  been  correctly  placed  in  Eukraiohelea:  E.  africana  Ingram 
and  Macfie,  1921,  E.  versicolor  Ingram  and  Macfie,  1921,  and  E.foyi 
Ingram  and  Macfie,  1922,  all  from  West  Africa;  Stilobezzia  (Eukraio- 
helea) aberrans  Johannsen,  1931,  from  Java;  S.  (E.)  esakiana  Tokunaga, 
1940,  from  the  Caroline  Islands;  and  Eukraiohelea  amnigena  Macfie, 
1935,  from   Brazil.     Eukraiohelea  inusitata  Johannsen,    1946,    from 


62  PROCEEDINGS    OF    THE    NATIONAL    MUSEUM  vol.  103 

Guam  is  apparently  a  synonym  of  esakiana  Tokunaga.  Palpomyia 
dorsofasciata  Lutz,  1914,  from  Brazil  was  correctly  referred  to  Eukraio- 
helea  by  Macfie  (1935).  Parabezzia  poikiloptera  Ingram  and  Macfie, 
1922,  from  West  Africa  should  be  referred  to  Eukraiohelea  as  tenta- 
tively suggested  by  Johannsen  (1946),  but  the  position  of  S.  ugandae 
Ingram  and  JMacfie,  1923,  from  East  Africa,  which  Johannsen  also 
believed  to  belong  here,  is  more  doubtful.  De  Meillon  (1938),  more- 
over, states  that  versicolor  and  poikiloptera  should  be  placed  in  the 
subgenus  Stilobezzia,  since  material  he  determined  as  these  two  species 
from  Lourengo  Marques,  Mozambique,  possessed  a  small  but  distinct 
first  radial  cell. 

1.  Stilobezzia  (Eukraiohelea)  elegantula  (Johannsen),  1908,  new  combination 

Figure  II,  g 

Bezzia  elegantulus  Johannsen,  Kansas  Univ.  Sci.  Bull.,  vol.  4,  p.  109,  1907  ( 9  , 

Kansas) . 
Probezzia  elegantula,  Malloch,  Proc.  Biol.  Soc.  Washington,  vol.  27,  p.  137,  1914. 
Parabezzia  elegantula,  Malloch,  Bull.  Illinois  State  Lab.  Nat.  Hist.,  vol.  10,  p.  359, 

1915. — Johannsen,  Journ.  New  York  Ent.  Soc,  vol.  42,  p.  345,  1934. 
Eukraiohelea  elegantula,  Johannsen,  Ann.  Ent.  Soc.  Amer.,  vol.  36,  p.  781,  1943. 

Description. — Female:  Length  2.0  mm.,  wing  1.5  mm.  by  0.7  mm. 
A  yellowish  species,  some  specimens  with  a  greenish  tinge;  antennae 
brown,  pedicel  and  narrow  bases  of  flagellar  segments  yellow;  palpi 
dark  brown.  Mesonotum  deep  yellow,  with  broad  sublateral  brown 
bands;  scutellum  yellow  in  center,  ends  brown;  postscutellum  brown; 
pleura  grayish,  a  large  black  spot  on  mesopleuron.  Legs  yellow;  coxae 
dark  below;  trochanters  brown;  apex  of  hind  femur,  base  of  hind  tibia, 
apices  of  all  tibiae  and  narrow  apices  of  all  tarsal  segments  brown. 
Wings,  including  veins,  grayish  hyaline ;  halteres  yellow,  sides  of  knobs 
black,  flat  end  of  knobs  pale  green.  Abdomen  dark  grayish  brown, 
apices  of  tergites  with  narrow  white  bands. 

Antennae  very  long  and  slender.  Mesonotum  with  long  black 
hairs  in  rows;  scutellum  with  four  long  black  marginal  bristles  and  a 
few  short  hairs.  Legs  with  hairs  fine  except  on  hind  tibia  where  they 
are  long  and  black ;  proportions  of  segments  of  hind  leg  as  in  table  1 ; 
fore  femur  with  two  or  three  small  black  spines  midway  on  inner 
margin;  basitarsus  with  a  long  black  basal  and  a  smaller  distal  spine 
on  midlegs  and  hind  legs;  fifth  tarsal  segment  with  two  pairs  of  long 
black  ventral  batonnets;  claws  slender  and  unequal,  the  outer  claw 
as  long  as  fifth  segment,  the  other  about  a  third  as  long.  Wing  with- 
out macro trichiae;  first  radial  cell  absent,  r-m  crossvein  and  Ri  nearly 
forming  an  oblique  line,  lengths  of  the  two  sections  of  costa  in  ratio 
of  0:6;  petiole  of  media  as  long  as  r-m  crossvein. 


HELEID    MIDGES,    GENUS    STILOBEZZIA — ^WIRTH  63 

Male:  Ninth  sternite  nearly  four  times  as  broad  as  long,  with  deep 
quadrate  emargination  three-fourths  way  to  base  on  mesal  third,  with 
spiculate  margms,  the  posterior  membrane  bare;  ninth  tergite  rounded, 
with  a  pair  of  very  small  iingerlike  setigerous  apicolateral  lobes. 
Basistyles  simple,  about  twice  as  long  as  broad;  dististyles  about  two- 
thirds  as  long  as  basistyles,  very  broad  with  truncate  tips.  Aedeagus 
with  an  oblique  pair  of  rather  stout,  slightly  sinuate,  sclerotized  bars 
with  pointed  apices;  an  irregular  hyalme  membranous  lobe  from  pos- 
terior margins.  Parameres  very  long,  slender  and  rodlike,  with  slender 
lateral  apodemes;  stems  about  a  fourth  again  as  long  as  basistjdes, 
subparallel,  the  apices  slightly  swollen,  palplike,  and  bent  ventro- 
laterad. 

Type. — In  Snow  collection.  University  of  Kansas,  9,  Kansas, 
Douglas  County,  Lawrence. 

Material  examined. — Florida:  Crystal  River,  Citrus  County,  July 
7,  1949,  Sept.  18,  1950,  Hudson,  2  99,  Miami,  Dade  County,  Oct.  15, 
1947,  Bm-en,  1  9;  Panama  City  Beach,  Bay  County,  July  2,  1950, 
McElvey,  1  cf 

Louisiana:  Baton  Rouge,  Apr.  30,  1947,  Wirth,  1  9. 

Jamaica:  Newport,  Feb.  22,  1937,  Chapin  and  Blackwelder,  1  9. 

Remarks. — S.  (E.)  africana  Ingram  and  Macfie,  from  West  Africa, 
the  genotype  of  Eukraiohelea,  is  almost  identical  with  elegantula,  but 
is  smaller  and  the  male  genitalia  differ  markedly;  the  ninth  sternite 
does  not  have  the  mesal  excavation,  the  dististyles  have  the  apices 
slenderer  and  rounded,  the  apices  of  the  parameres  are  straight  and 
pointed,  and  the  membrane  of  the  aedeagus  is  prolonged  mesad  in  a 
sharp  cone. 

Subgenus  Neostilobezzia  Goetghebuer 

Neostilobezzia  Goetghebuer,  in  Lindner,  Die  Fliegen  der  Palaearktischen  Region, 
Lief.  78,  p.  53,  1934.  (Genotype,  Ceratopogon  ochraceus  Winnertz,  1852, 
by  present  designation.) 

Discussion. — Goetghebuer  (1934)  distinguished  this  subgenus  from 
Stilohezzia  by  the  presence  of  macrotrichiae  at  the  end  of  the  wing, 
the  wing  of  Stilohezzia  being  entirely  bare.  In  addition,  all  the  species 
known  to  me  are  yellowish  or  brown  in  ground  color,  varying  from 
almost  whitish  to  almost  black,  and  the  male  genitalia  have  the 
aedeagal  sclerites  decidedly  bent  or  humped  in  the  middle  and  the 
basistyle  lacks  the  inner  lobe.  Species  with  these  characters  are 
widely  distributed  around  the  world.  It  is  especially  significant  that 
most  of  the  species  described  from  the  temperate  regions  belong  to 
this  subgenus,  while  the  tropical  species  are  predominantly  the  sub- 
genus Stilohezzia. 


64  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  103 

2.  Stilobezzia  {Neostilobezzia)  fuscula  Wirth,  1952 

Figure  11,  e 

Stilobezzia  fuscula  Wirth,  Univ.  California  Publ.  Ent.,  vol.  9,  p.  204,  1952  ( 9  , 
California) . 

Diagnostic  characters.— This  species  resembles  S.  (N).  lutea  (Mal- 
loch),  1918,  in  its  uniformly  yellowish  color,  unmarked  wings  with 
macrotrichiae,  and  mesonotum  with  indistinctly  darker  vittae;  but 
differs  in  being  somewhat  duller  and  darker,  with  dark  antennal 
pedicel  and  halteres,  and  in  having  the  body  hairs  much  softer. 

Description. — Female:  Length  1.4  mm.,  wing  1.5  mm.  by  0.5  mm. 
Head  grayish  brown  pruinose,  pedicel  of  antennae  and  palpi  brown, 
antennal  flagellum  yellowish  brown;  segments  in  proportion  of 
20 :  12 :  12 :  12 :  12 :  12 :  12 :  12 :  20 :  20 :20 :  20 :  30.  Palpal  segments  m  pro- 
portion of  4:8:15:10:10,  third  segment  with  three  long,  stalked  sen- 
sillae. 

Mesonotimi  dull  yellowish  brown,  with  coarse  pruinosity,  a  narrow 
median  longitudinal  vitta  and  a  broad  patch  on  each  side  from  humeral 
pits  to  wing  base  darker  brown;  most  of  bristles  removed,  but  those 
few  remaining  long  and  dark  brown.  Scutellum  bright  yellow  in 
middle,  brownish  on  sides,  apparently  with  four  marginal  bristles. 
Postscutellum  and  plem'a  yellowish  brown  pruinose.  Legs  dull  dark 
yellowish;  hairs  short,  except  row  of  long  bristles  on  outer  edge  of  hind 
tibia;  inner  claw  as  long  as  fifth  tarsal  segment. 

Wings  grayish  hyaline,  anterior  veins  brownish;  second  anterior 
radial  cell  about  five  times  as  long  as  fu'st;  macrotrichiae  thick  at  apex 
of  cell  R5  and  along  and  beyond  intercalary  fork,  a  few  also  along  tip 
of  veins  Mi  and  M2  and  at  apex  of  cell  Mi.  Halteres  brown,  flat  end  of 
knob  dull  yellowish. 

Abdomen  dull  dark  brown,  a  patch  of  long  brown  hairs  on  sides 
of  first  segment.  Spermathecae  two,  oval,  with  ducts  sclerotized 
a  short  distance;  a  rudimentary  third  spermatheca  present. 

Male:  Similar  to  the  female  but  with  the  usual  sexual  differences; 
antennal  plumes  hght  brown;  thorax  and  abdomen  infuscated  light 
brown;  scutellum  yellow. 

Ninth  sternite  about  three  times  as  broad  as  long,  with  mesal 
posterior  excavation  halfway  to  base,  posterior  membrane  bare;  ninth 
tergite  rather  long  and  rounded,  with  a  pair  of  long,  rounded,  seti- 
gerous,  apicolateral  lobes.  Basistyle  nearly  three  times  as  long  as 
broad,  with  a  black  spine  borne  on  a  small  lobe  with  a  long  fine  hau- 
about  midway  of  mesal  margin;  dististyle  as  long  as  basistyle,  slender 
and  slightly  curved  to  apex.  Aedeagus  with  a  pair  of  oblique,  lateral, 
sclerotized  bars  with  median  apices  forked,  the  inner  arm  in  line 
with  base  and  longer  than  the  other  arm.     Parameres  with  slender 


HELEID    MIDGES,    GENUS    STILOBEZZIA — WIRTH  65 

basal  apodemes,  stems  long  and  slender,  with  bases  slightly  swollen, 
each  with  apex  gradually  narrowed  to  a  fine  apical  filament  abruptly 
bent  and  recurved  on  distal  fourth. 

Type.—VSNM  59946,  9,  California,  Tulare  County,  6  miles  east  of 
Orosi.  (In  the  original  publication  the  sex  was  erroneously  given 
as  cT.) 

Material  examined.— Caliiormsi:  6  miles  east  of  Orosi,  Tulare 
County,  July  8,  1947,  Wirth,  2  99  (type  and  paratype). 

Utah:  Moab,  June  8,  1948,  Knowlton,  Harmston,  and  Wood,  at 
light.  Id". 

Remarks. — The  male  genitalia  of  S.fuscula  are  practically  identical 
with  those  of  S.  macfiei  Lane,  1947,  from  Brazil.  However,  the  latter 
species  has  the  mesonotum  polished  black,  with  lighter  areas,  and 
under  certain  light  a  spot  on  each  side  before  wing  base  has  purplish 
reflections.  Moreover,  the  absence  of  macrotrichiae  on  the  wings 
would  place  S.  macfiei  in  the  subgenus  Stilobezzia. 

3.  Stilobezzia  {Neostilohezzid)  lutea  (Malloch),  1918 

Figure  11,  6 

Hartomyia  gilva  Malloch,  not  Coquillett  (misidentification) ,  Bull.  Illinois  State 

Lab.  Nat.  Hist.,  vol.  10,  p.  343,  1915. 
Hartomyia  lutea  Malloch,  Bull.  Brooklyn  Ent.  Soc,  vol.  13,  p.  18,  1918,  (  9  ,  Illinois). 
Stilobezzia  lutea,  Johannsen,  Ann.  Ent.  Soc.  Amer.  vol.  36,  p.  781,  1943. 
Stilobezzia  mallochinoSmSin,  Ent.  ^ews,  vol.  Z5,  p.  2SS,  1924,  (c?,    9;  New  York, 

Pennsylvania). — Johannsen,  Ann.  Ent.  Soc.  Amer.,    vol.   36,   p.    781,    1943 

(eastern  United  States).     (New  synonymy.) 

Description. — Female:  Length  2.5  mm.,  wing  1.9  mm.  by  0.7  mm. 
Yellow,  including  anterior  wing  veins,  antennae,  and  legs ;  mesonotum, 
except  humeri,  and  apex  of  abdomen  rufous  brown.  Vestiture  of 
very  long  coarse  brown  bristles,  those  on  scutellum  as  long  as  breadth 
of  scutellum. 

Male:  Ninth  sternite  nearly  three  times  as  broad  as  long,  with  a 
very  broad,  shallow,  posterior  emargination,  the  membrane  spiculate; 
ninth  tergite  conical,  with  a  prominent,  submedian  pair  of  long,  fleshy, 
setose  lobes.  Basistyles  simple,  twice  as  long  as  broad;  dististyles 
slightly  longer  than  basistyles,  curved,  and  gradually  attenuated, 
with  pointed  apices.  Aedeagus  with  a  pair  of  oblique  sinuate 
sclerites,  bases  of  which  are  articulated  with  ventral  root  of  basistyle, 
apices  close  together,  pointed,  and  bent  ventrad.  Parameres  with 
stout  basal  apodemes,  the  stems  close  together  and  slightly  sinuate, 
flattened  and  slightly  expanded  past  middle,  with  apices  pointed  and 
somewhat  curved  ventrad. 

Type. — In  collection  Illinois  Natural  History  Survey,  9,  Illinois, 
Elizabeth. 

Material  examined. — New  Hampshire:  Center  Harbor,  July  9, 
Dyar,  1  9;  Franconia,  Slosson,  1  cT. 

232991—53 2 


66  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  103 

Connecticut:   Green  Falls,  June  31,  1935,  Chapman,  1  cf ,  1  9. 

Rhode  Island:  Westerly,  June  10,  1936,  Chapman,  1  <^. 

New  York:  Armonk,  Bemus  Pomt,  Canadarago  Lake,  Kast 
Bridge,  Millwood,  Poughkeepsie,  Rome,  June,  July  1934-36,  H.  K. 
Townes,  12  cfd",  7  99. 

New  Jersey:  Moorestown,  June  6,  1936,  H.  and  M.  Townes,  1  9; 
Riverside,  June  18,  1939,  H.  K.  Townes,  1  cf . 

Delaware:  Smyrna,  June  8,  1937,  Bradley,  1  cf. 

Maryland:  Cabin  John,  June  18,  1920,  Aldrich,  1  9;  Glen  Echo, 
July  1,  1923,  Malloch,  1  9;  Plummer's  Island,  June  18,  1914,  Shannon, 
1  9. 

District  of  Columbia:  Rock  Creek  Park,  June  17,  1920,  Aldrich,  1  cf. 

Virginia;  Dead  Run,  June  6,  1914,  Shannon,  1  9;  Falls  Church, 
June  7,  1914,  Shannon,  2  99,  and  July  4,  8,  1950,  Wirth,  4  cT"  cT,  7  99; 
Glencarlyn,  June  7,  1935,  Malloch,  1  cf ,  1  9. 

South  Carolina:   Greenville,  June  4,  1933,  H.  K.  Townes,  1  cf. 

Indiana:  Lafayette,  June  26,  1916,  Aldrich,  1  cf ,  4  99. 

Illinois:  Elizabeth,  July  7,  1917,  Malloch,  1  9  (type  of  Hartomyia 
lutea  Malloch), 

Michigan:    Cheboygan  County,  July  17,  1942,  Sabrosky,  1  9. 

Iowa:  Maquoka  Caves  State  Park,  Jackson  Count}'',  Jul}'  3,  1949, 
Laffoon,  1  9;  Pikes  Peak  State  Park,  Clayton  County,  July  4,  1949, 
Laffoon,  1  9;  Sioux  City,  June,  July  1949-50,  Slater  and  Laffoon, 
6  d'c?',  11  99. 

Remarks. — Alalloch  had  misidentified  the  male  of  this  species  as 
Hartomyia  gilva  (Coquillett),  which  is  an  Atrichopogon,  in  1915,  and 
in  1918  described  the  female  of  the  same  species  as  Hartomyia  lutea, 
new  species.  Hoffman  discovered  that  Malloch's  determination  of 
gilva  was  incorrect  and  described  the  species  as  Siilohezzia  mallochi, 
1924,  in  part  from  Malloch's  material.  Since  lioffman  made  no 
mention  of  Malloch's  description  of  hitea,  he  ma}^  have  been  unaware 
of  it,  or  he  may  not  have  been  able  to  recognize  Malloch's  species. 
The  present  synonymy  has  been  made  after  an  examination  of  the 
type  of  lutea  and  specimens  determined  as  gilva  by  IVfalloch  in  the 
Illinois  Natural  History  Survey  through  the  kindness  of  Dr.  H.  H. 
Ross,  and  of  specimens  in  the  U.  S.  National  Museum  determined  by 
Hoffman  as  mallochi.  Hoffman's  detailed  description  of  this  species 
needs  no  elaboration  except  for  the  characters  of  the  male  genitalia. 

4.  Stilobezzia  {Neostilobezzia)  stonei,  new  species 

Figure  11,  c 

Diagnostic  characters. — A  large  brownish  species;  mesonotum  and 
scutellum  dull  yellowish ;  legs  yellowish;  midfemora,  hind  femora,  and 
tibiae  dark  brown;  wings  grayish  brown;  hal teres  dull  yellow. 


HELEID    MIDGES,    GENUS    STILOBEZZIA — ^WIRTH  67 

Description. — Female:  Length  1.8  mm.,  wing  1.8  mm.  by  0.7  mm. 
Head  dull  brownish,  including  antennae  and  palpi,  with  a  few  long 
black  hairs.  Antenna  as  long  as  head  and  thorax  combined;  flagellar 
segments  in  proportion  of  20:15:15:15:15:16:17:18:25:25:25:25:30. 
Palpal  segments  slender,  in  proportion  of  5:10:12:6:10,  third  segment 
slightly  swollen  on  basal  two-thirds,  with  a  small  pit  at  distal  fourth. 

Thorax  dull  brown,  mesonotum  and  scutellum  dull  yellowish, 
postscutellum  and  pleura  dark  brown.  Mesonotum  with  moderate 
to  long  brown  hairs  in  rows;  scutellum  with  about  ten  long  brown 
marginal  hairs.  Legs  dull  brown,  usually  all  of  forelegs,  midtarsus 
and  hind  tarsus  yellowish.  Proportions  of  segments  of  hind  leg  as  in 
table  1;  legs  moderately  slender,  hairs  fine,  not  very  long  except 
those  on  hind  tibia  about  twice  the  diameter  of  segment;  basi tarsi 
with  ventral  spines,  three  or  four  on  basal  half  on  forelegs  and  midlegs, 
one  at  base  on  hind  leg;  claws  unequal,  rather  strong  and  curved,  the 
longer  nearly  as  long  as  fifth  segment,  the  other  about  half  as  long. 

Wings  evenly  infuscated,  the  veins  slightly  darker,  a  few  macro- 
trichiae  at  apices  of  cells  Rg  and  Mi.  First  radial  cell  large,  about  a 
third  the  length  of  second;  costa  extending  to  0.8  wing  length;  cross- 
vein  r-m  faint  in  middle,  oblique;  petiole  of  media  as  long  as  crossvein. 
Halteres  yellow.  Abdomen  dark  brown  with  short  fine  hairs;  two 
large,  slightly  unequal  ovoid  spermathecae  and  a  vestigial  third 
present,  ducts  not  sclerotized. 

Male:  Similar  to  the  female  but  with  the  usual  sexual  differences; 
antennal  plumes  brownish,  knees  quite  dark.  Ninth  sternite  about 
three  times  as  broad  as  long,  with  broad  posterior  excavation  more 
than  halfway  to  base,  the  membrane  bare;  ninth  tergite  rounded,  with 
small,  submedian,  setose,  apicolateral  lobes.  Basistyles  simple,  twice 
as  long  as  broad ;  dististyles  0.9  times  as  long  as  basistyles,  slender  and 
nearly  straight,  their  apices  slightly  knobbed.  Aedeagus  with  an 
oblique  pair  of  slightly  curved  sclerites  with  distal  ends  close  together, 
sharp-pointed,  and  bent  ventrad;  apparently  bearing  a  pair  of  hyaline 
membranous  lobes  on  posterior  margins  and  attached  to  membrane 
connecting  dorsal  roots  of  basistyles  on  ventral  side  of  parameres. 
Parameres  with  broad,  platelike,  basal  apodemes;  stems  very  broad 
and  flattened,  gradually  expanded  distad  with  a  short  lateral  point  and 
a  longer,  slenderer,  ventrally  bent,  distal  point,  their  distal  margins 
obliquely  truncated. 

Types.— VSNM  60964,  holotype,  d",  and  aflotype,  Falls  Church, 
Va.,  July  8,  1950,  Wirth.  Paratypes:  Virginia:  16  cTcf ,  24  99,  same 
data  as  type.  Georgia:  2  cf  c?",  17  99,  Thomasville,  May  1949,  Palmer. 
Florida:  2  99,  Innerarity  Point,  May  4,  1950,  Eathert;  1  9,  Panama 
City  Beach,  May  6,  1949,  McElvey. 


68  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  103 

Remarks. — Related  to  S.  lutea  (Malloch)  in  general  features,  but 
readily  distinguished  by  the  more  extensive  brown  coloration, 
finer  vestiture  and  the  characteristic  broad,  obliquely  truncated,  two- 
pointed,  platelike,  male  parameres.  This  species  is  named  in  honor  of 
Dr.  Alan  Stone. 

Subgenus  Stilobezzia  Kieffer,  1911 

Stilobezzia  KiefiFer,  Rec.  Indian  Mus.,  vol.  6,  p.  118,  1911,  and  vol.  9,  p.  184,  1913; 
Bull.  Ent.  Soc.  France,  p.  192,  1919. — Goetghebuer,  Mem.  Mus.  Hist.  Nat. 
Belgique,  vol.  8,  p.  59,  1920. — Carter,  Ingram,  and  Macfie,  Ann.  Trop.  Med. 
Parasit.,  vol.  15,  p.  324,  1921. — Edwards,  Trans.  Ent.  Soc.  London,  vol.  74, 
p.  411,  1926;  Notulae  Ent.,  vol.  9,  p.  9,  1929.— Johannsen,  Arch.  Hydrobiol., 
SuppL,  vol.  9,  p.  430,  1931;  Ann.  Ent.  Soc.  Amer.,  vol.  36,  p.  781,  1943.— 
Ingram  and  Macfie,  The  Diptera  of  Patagonia  and  southern  Chile  .  .  ., 
pt.  2,  fasc.  4,  p.  191,  1931. — de  Meillon,  Proc.  Ent.  Soc.  London,  ser.  B,  vol. 
7,  p.  266. — Tokunaga,  Philippine  Journ.  Sci.,  vol.  72,  p.  155,  1940. — Lane, 
Rev.  Ent.,  vol.  18,  p.  197,  1947.— Lee,  Proc.  Linn.  Soc.  New  South  Wales, 
vol.  72,  p.  345,  1948.— Wirth,  Univ.  California  Publ.  Ent.,  vol.  9,  p.  202,  1952. 
(Genotype,  Ceratopogon  notatus  de  Meijere,  as  Stilobezzia  f estiva  KiefiFer; 
original  designation.) 

Hartomxjia  Malloch,  Bull.  Illinois  State  Lab.  Nat,  Hist.,  vol.  10,  p.  339,  1915. 
(Genotype,  Ceratopogon  pictus  Coquillett;  original  designation.) 

Diagnosis. — Two  radial  cells  present;  wing  without  macrotrichiae 
at  apex. 

5.  Stilobezzia  (Stilobezzia)  antennalis  (Coquillett),  1901 

Figure  12,  e 

Ceratopogon  antennalis  Coquillett,  Proc.  U.  S.  Nat.  Mus.,  vol.  23,  p.  606,  1901 

(  9  ,  District  of  Columbia). 
Ceratolophiis  antennalis,  Kieffer,  Genera  insectorum,  fasc.  42,  p.  60,  1906. 
Johannseniella  antennalis,  Malloch,  Bull.  Illinois  State  Lab.  Nat.  Hist.,  vol.  10, 

p.  227,  1914. 
Hartomyia  antennalis,  Malloch,  Bull.  Illinois  State  Lab.  Nat.  Hist.,  vol.  10,  p.  343, 

1915  (cf  ;  Illinois,  Indiana). 
Stilobezzia  antennalis,  Johannsen,  Ann.  Ent.  Soc.  Amer.,  36:  781,  1943. 

Description. — Female:  Length  1.5  mm.,  wing  1.6  mm.  by  0.6  mm. 
Head,  thorax,  and  abdomen  polished  black;  antennae  and  legs  yellow, 
midfemur  and  hind  femur  black  except  at  extreme  bases.  Wings  bare 
and  hyaline,  anterior  veins  yellowish,  knob  of  hal teres  black.  Palpi 
and  last  five  flagellar  segments  slightly  infuscated,  flagellar  segments 
in  proportion  of  18:11:11:11:11:11:11:15:25:25:25:25:30.  Palpal 
segments  in  proportion  of  4:8:14:8:15,  third  segment  not  swollen, 
pit  absent.  Proportions  of  segments  of  hind  leg  as  in  table  1 ;  basitarsi 
without  spines,  fifth  segment  with  a  pair  of  long  ventral  batonnets; 
claws  long,  simple,  and  very  unequal,  the  short  claw  fused  with  base 
of  and  about  a  third  as  long  as  the  other.  First  radial  cell  about  a 
fourth  as  long  as  second;  r-m  crossvein  vertical,  petiole  of  media  to  a 


HELEID    MIDGES,    GENUS    STILOBEZZIA — WIRTH  69 

third  the  length  of  Mi.  Spermathecae  two,  the  larger  oval,  the 
smaller  subspherical  and  a  third  the  diameter  of  other,  a  minute 
vestigial  third  present. 

Male:  Similar  to  the  female  but  with  the  usual  sexual  differences; 
plumes  of  antennae  brown.  Ninth  sternite  a  narrow  transverse  ante- 
rior band,  the  membrane  spiculate;  ninth  tergite  evenly  rounded 
caudad,  the  apicolateral  lobes  not  prominent.  Basis tyle  with  large 
irregular  lobe  on  inner  side  to  distal  third;  dististyle  rather  strongly 
bent  past  middle  and  tapered  to  pointed  tip.  Aedeagus  with  a  pair 
of  oblique,  straight,  sclerotized  bars  with  distal  apices  scarcely  meet- 
ing. Parameres  a  pair  of  strongly  sclerotized,  large,  strongly  sinuate 
rods  with  pointed  apices  and  large  basal  knobs. 

Type.—USNM  5481,  9,  District  of  Columbia. 

Material  examined. — New  York:  Ithaca,  June  11,  1935,  H.  K. 
Townes,  1  9;  Rome,  June  24,  1934,  H.  K.  Townes,  1  d'. 

New  Jersey:  Moorestown,  June  6,  1939,  H.  and  M.  Townes,  1    c?. 

Maryland:  Dorchester,  July  10,  1907,  Barber,  1  9;  Mayo,  July  22, 
1950,  Wirth,  2  cf  cf,  1  9;  Plummer's  Island,  June  30,  1914,  Shannon, 

1  9,  and  Aug.  18,  1913,  Viereck,  1  9. 

District  of  Columbia:  June,  1  9  (type) ;  Benning's,  July  21,  Aldrich, 

2  99. 

Virginia:  June  IG,  1903,  Pergande,  2  cfcf,  3  99;  Dead  Run,  June 
22,  1915,  Shannon,  1  9,  Falls  Church,  July  4,  8,  1950,  Wu'th,  3  c^  d', 
4  99. 

Michigan:  Midland  County,  July  17,  1944,  Dreisbach,  1  9;  Traverse 
City,  June  17,  1943,  Sabrosky,  1  cf. 

lUinois:  Freeport,  July  4,  1917,  Malloch,  3  cf  cf;  Galena  Junction, 
July  8,  1917,  Malloch,  1  9;  White  Heath,  June  4,  1939,  Dirks,  1  9. 

Tennessee:  Ivnoxville,  May  25,  Aldrich,  1  9. 

Mississippi:  West  Point,  Aug.  11,  1904,  Barber,  1  cf. 

Louisiana:  Baton  Rouge,  May  4,  16,  1947,  Wirth,  4  cf  cf,  8  99. 

6.  Stilobezzia  {StUobezsia')  beckae,  new  species 

Figure  11,  / 

Diagnostic  characters. — A  large,  dark  brown  species  with  silvery 
pruinose  pattern  on  mesonotum  and  abdomen;  wing  with  two  darker 
and  one  lighter  spots  on  anterior  margin;  knobs  of  hal teres  black; 
mesonotum  with  strong  anterior  tubercle;  fifth  tarsal  segment 
unspined. 

Description. — Female:  Length  2.5  mm.,  wing  1.8  mm.  b}^  0.7  mm. 
Head  black  with  dense  whitish  pruinosity,  antennae  brown,  bases  of 
flagellar  segments  3  to  10  yellowish;  palpi  dark  brown.  Flagellar 
segments  of  antenna  in  proportion  of  16:12:15:20:20:20:20:22:28: 


70  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  103 

30:32:32:42.  Palpal  segments  in  proportion  of  5:12:15:8:12, 
third  segment  slightly  swollen,  with  large,  shallow  sensory  pit  near 
apex. 

Thorax  dark  reddish  brown,  with  dense  silvery  pruinescence  in 
irregular  patches;  mesonotum  long,  with  prominent,  conical,  anterior 
spine,  the  anterior  portion  markedly  sloping,  posterior  portion  flat- 
tened between  wing  bases;  covered  with  sparse,  long,  brown  hairs; 
scutellum  with  four  long,  fine,  brown  hairs.  Legs  dark  brown,  all 
femora  and  fore  tibiae  with  obscure  broad  median  light  bands,  mid- 
tibia  and  hind  tibia  with  narrow  sub-basal  light  rings,  tarsi  yellow 
except  fifth  segment  of  each  brown.  Proportions  of  segments  of 
hind  leg  as  in  table  1 ;  legs  with  moderately  long,  black  hairs,  wliich  are 
stronger  on  hind  tibia ;  basitarsi  with  three  ventral  spines  on  foreleg, 
four  on  midleg,  and  one  at  base  on  hind  leg;  fifth  segment  without 
ventral  batonnets;  claws  very  slender  and  slightly  unequal,  the  longer 
as  long  as  fifth  segment,  the  other  about  0.6  to  0.8  as  long. 

Wing  hyaline,  without  macrotrichiae,  veins  infuscated,  a  large 
brownish  spot  over  first  radial  cell  from  vein  Ri  to  crossvein  r-m; 
apex  of  second  radial  cell  with  bro^vn  spot,  then  a  short  white  area 
with  apex  of  wing  weakly  infuscated.  First  radial  cell  well  developed, 
one-fourth  as  long  as  second ;  r-m  crossvein  faint  in  middle,  forming  an 
oblique  line  with  Ri,  petiole  of  media  as  long  as  crossvein.  Knobs  of 
halteres  black. 

Abdomen  black,  distal  half  of  tergites  1  to  3  and  narrow  distal 
margins  of  remaining  tergites  with  silvery,  pruinose  bands.  Sper- 
mathecae  two,  subequal,  pyriform. 

Male:  Similar  to  the  female  but  with  the  usual  sexual  differences; 
antennal  plumes  golden.  Ninth  sternite  over  twice  as  broad  as  long, 
with  caudal  excavation  to  half  its  length,  the  membrane  bare;  ninth 
tergite  rounded  with  a  pair  of  membranous,  setigerous,  apicolateral 
lobes.  Basistyles  simple,  about  2.5  times  as  long  as  broad;  dististyles 
slender,  nearly  as  long  as  basistyles,  gently  curved,  with  pointed 
incurved  apices.  Aedeagus  with  a  pair  of  heavj^,  oblique,  crooked, 
lateral  sclerites  meeting  mesad  subapically,  with  apices  pointed  and 
abruptly  bent  ventrolaterad.  Parameres  with  heavy,  platelike, 
V-shaped,  basal  apodemes;  stems  straight  and  contiguous  mesad, 
slightly  expanded  midway,  with  lateral  wall  folded  ventrad  and  mesad 
and  apparently  fusing  on  midline  on  distal  third;  extreme  apices 
slender,  flattened  dorsoventrally  and  abruptly  bent  ventrad. 

Types. — USNM  60695,  holotype,  cf,  and  allotype.  Crystal  River, 
Citrus  County,  Fla.,  June  7,  1950,  Hudson,  light  trap.  Parat3T)es: 
Florida:  10  cf  cf ,  17  99,  same  data  as  type  except  dates,  June  5-July  17, 
1949-50;  6  99,  Grayton  Beach,  Walton  County,  May  10,  1949,  Butler; 
15  99,  Santa  Rosa,  Walton  County,  June  7,  1949,  Peterson.  Missis- 
sippi: 4  99,  Horn  Island,  June,  July,  1944,  Richmond. 


HELEID    MIDGES,    GENUS    STILOBEZZIA — ^WIRTH  71 

Additional  material  examined. — Peru:  Iquitos,  March,  April  1931, 
Shannon,  1  9. 

Remarks. — Closely  allied  to  S.  paulistensis  Lane,  1947,  from  Brazil 
which  differs,  however,  in  having  the  legs  yellowish  rather  than  black- 
ened, in  lacking  the  dark  spot  at  apex  of  second  radial  cell,  and  in 
having  the  lateral  sclerites  of  the  male  aedeagus  shorter,  with  apices 
capitate  and  bent  mesad  rather  than  slender  and  bent  laterad.  S. 
rabelloi  Lane,  1947,  is  also  closely  related,  but  is  a  larger,  hairier 
species  with  three  dark  wing  spots,  including  one  at  apex  of  first 
radial  cell,  and  has  several  pairs  of  batonnets  on  fifth  tarsal  segment. 
I  am  happy  to  name  this  species  for  Mrs.  Elisabeth  C.  Beck  of  Arling- 
ton, Fla.,  who  has  kindly  sent  me  so  many  interesting  Florida  heleids. 

7.  Stilobezzia  {Stilobezzia)  hicolor  Lane,  1947 

Figure  11,  i 
Stilobezzia  hicolor  Lane,  Rev.  Ent.,  vol.  18,  p.  208,  1947  (c^,  Brazil). 

Description. — Female:  (Here  described  for  the  first  time.)  Length 
1.5  mm.,  wing  1.5  mm.  b}^  0.6  mm.  Head  dark  brown,  antennal 
pedicel,  clypeus,  and  palpi  yellowish.  Thorax  and  abdomen  polished 
black,  with  metallic  bluish  violet  reflections.  Mesonotum  rather 
broad  and  flat,  with  sparse,  long,  brown  hairs,  scutellum  with  four 
marginal  bristles.  Alidcoxa  and  hind  coxa  black,  rest  of  legs  yellow, 
except  knees  and  fifth  tarsal  segments  dark.  Legs  with  fine  hairs; 
proportions  of  segments  as  in  table  1 ;  basitarsi  unspined ;  fifth  segment 
with  a  pair  of  long  black  batonnets  at  base:  claws  slender  and  unequal, 
the  outer  claw  as  long  as  fifth  segment,  the  inner  about  a  third  as  long. 
Wing  hyaline,  with  violet  reflections,  anterior  veins  yellow;  macro tri- 
chiae  absent.  First  radial  cell  distinct,  a  sixth  as  long  as  second; 
petiole  of  media  about  twice  as  long  as  crossvein  r-m.  Halteres  black. 
Abdomen  convex  above,  somewhat  petiolate,  with  a  row  of  long,  brown 
hairs  across  each  tergite. 

Male:  As  in  the  female  but  with  the  usual  sexual  differences; 
plumes  of  antennae  brown.  Ninth  sternite  a  narrow  anterior  band, 
the  posterior  membrane  spiculate;  ninth  tergite  rounded,  with  incon- 
spicuous, rounded,  setigerous,  apicolateral  lobes.  Basistyle  with  a 
small  lobe  at  half  the  length  of  inner  margin;  distist3de  about  as  long 
as  basistyle,  tapered  to  slender  tip.  Aedeagus  with  an  only  slightly 
oblique  pair  of  sclerotized  bars,  stout  in  the  Texas  specimen  figured, 
but  slenderer  in  the  tropical  specimens.  Parameres  with  lateral 
apodemes  bent  caudad;  stems  straight,  stout  and  rodlike,  with  beak- 
like apices  abruptly  bent  laterad. 

Type. — In  coUection  University  of  Sao  Paulo,  Brazil,  cf ,  Brazil, 
Sao  Paulo,  Osasco. 


72 


PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  103 


g.   ELEGANTULA 


h.  COQUILLETTI 


I.  BICOLOR 


Figure  11. — Male  genitalia  of  Stilobezzia  species:  a,  S.  (S.)  rabelloi  Lane;  b,  S.  (N.)  lutea 
(Malloch);  c.  S.  {N.)  stonei,  new  species;  d,  S.  (S.)  punctipes,  new  species;  e,  S.  {N.) 
fuscula  Wirth;/,  S.  (S.)  beckae,  new  species;  g,  S.  (£.)  elegantula  (Johannsen);  h,  S.  (S.) 
coquilletti  Kieffer;  i,  S.  (S.)  bicolor  Lane. 


HELEID  MIDGES,   GENUS  STILOBEZZIAWIRTH 


73 


a.    THOMSENAE 


b.    BULLA 


c.    SYBLEAE 


d,  VIRIOIS 


e.     ANTENNALIS 


/.     DIVERSA 


g.      PRUINOSA 


h.      GLAUCA 


Figure  12. — Male  genitalia  of  Stilobezzia  species:  a,  S.  (S.)  thomsenae,  new  species;  b,  S. 
(S.)  bulla  Thomsen;  c,  S.  (S.)  sybleae,  new  species;  d,  S.  (S.)  viridis  (Coquillett);  e,  S.  (S.) 
antennalis  (Coquillett);  /,  S.  (S.)  diversa  (Coquillett);  g,  S.  (S.)  pruinosa  Wirth;  h,  S. 
(S.)  glauca  Macfie. 


74  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  103 

Material  examined. — Texas:  San  Antonio,  Apr.  8,  1907.  Pratt,  1  cf . 
Costa  Rica:  Higuito,  Sdiild,  2  d'  d',  10  99. 

Puerto  Rico:  Anasco,  Apr.  22,  1933,  Faxon,  Mills,  and  Anderson, 
1  cf ,  4  99.     Bayamon,  Jan.  28,  1934,  Lesesner  and  Anderson,  1  9. 
Panamd:  Summit,  Canal  Zone,  November  1946,  Krauss,  2  cT  cf ,  1  9. 

8.  Stilohezzia  {Stilobezzia)  bulla  Thomsen,  1935 

Figure  12,  & 

Stilohezzia  bulla  Thomsen,  Journ.  New  York  Ent.  Soc,  vol.  43,  p.  289,   1935 
(cf,  9  ;  New  York);  Johannsen,  Ann.  Ent.  Soc.  Amer.,  vol.  36,  p.  781,  1943. 

Description. — Female.  Length  1.0  mm.,  wing  0.8  mm.  by  0.4  mm. 
Pruinose  grayish  brown;  scutellum  and  legs  yellowish ;hal teres  white; 
wings  clear,  veins  stramineous.  Antennal  segments  in  proportion  of 
12:8:9:10:10:10:10:10:13:15:15:15:20.  Palpi  short  and  slender, 
segments  in  proportion  of  5:7:7:5:7,  third  segment  not  swollen,  pit 
absent.  Basi tarsi  and  fifth  segment  without  spines;  claws  simple, 
slender  and  unequal,  the  longer  as  long  as  fifth  segment,  the  other 
a  third  as  long.  Wings  clear,  macrotrichiae  absent.  Spermathecae 
two,  oval,  slightly  unequal,  with  ducts  sclerotized  a  very  short 
distance. 

Male:  Ninth  sternite  a  narrow  anterior  ribbon,  the  membrane 
spiculate;  ninth  tergite  tapered  to  a  pair  of  prominent,  slender,  setose, 
apicolateral  lobes,  truncate  between  then*  bases.  Basistyle  slender, 
with  a  prominent,  heavily  sclerotized,  platelike  lobe  articulating  with 
the  median  sclerite  (?  of  aedeagus);  dististyle  about  half  as  long  as 
basistyle,  rather  stout  and  nearly  straight,  tip  bluntly  pointed,  with 
a  distinct  subapical  tooth  on  ventral  face  of  outer  edge.  Aedeagus 
with  basal  sclerites  reduced  to  very  small,  narrow  bars,  connected  at 
right  angles  by  a  pair  of  short,  barlike  sclerites  to  a  more  dorsal,  large, 
heavily  sclerotized,  transverse  sclerite  shaped  like  a  pair  of  buffalo 
horns,  with  points  articulating  with  inner  lobes  of  basistyles.  Para- 
meres  with  slender,  hook-shaped,  basal  apodemes,  stems  rodlike, 
slender  and  nearly  straight,  gradually  tapered  to  long,  simple,  fine 
tips  reaching  nearly  to  apices  of  lobes  of  ninth  tergite. 

Types. — In  collection  Cornell  University,  holotype,  cf ,  and  allo- 
type. New  York,  Ithaca,  McLean  Bogs. 

Material  examined. — Maryland:  Dorchester,  July  10,  1907,  Barber, 
1  d^. 

District  of  Columbia:  Bennings,  July  21,  Aldrich,  1  9. 

Virginia:  Falls  Church,  July  4,  8,  Aug.  6,  1950,  Wirth,  18  c^cf, 
24  99. 

Georgia:  Thomasville,  May  15-30,  1949,  Palmer,  1  9. 

Florida:  Crystal  River,  Sept.  18,  1950,  Hudson,  1  9. 


HELEID    MIDGES,    GENUS    STILOBEZZIA — WIRTH  75 

Louisiana:  Baton  Rouge,  Apr.  19,  1947,  Wirth,  1  cf ;  Kilbourne, 
May  10,  1947,  Wii-th,  1  9. 

9.  Stilobezzia  {Stilobezzia)  coquilletti  Kieffer,  1905 

Figure  11,  h 

Ceratopogon  pidus  Coquillett,  Journ.  New  York  Ent.  Soc,  vol.  13,  p.  60,  1905 

(9,  Virginia). 
Hartomyia  picta,  Malloch,  1915,  Bull.  Illinois  State  Lab.  Nat.  Hist.,  vol.  10,  p. 

341,  1915  (c?,  Illinois). 
Stilobezzia  coquilletti  Kieffer,  Ann.  Mas.  Nat.  Hungarici,  vol.  15,  p.  308,  1917 

(new  name  for  C.  picta  Coquillett  not  Meigen). — Johannsen,  Ann.  Ent.  Soc. 

Amer.,  vol.  36,  p.  781,  1943. 

Description. — Female:  Length  2.0  mm.,  wing  2.0  mm.  by  0.8  mm. 
Yellow,  in  some  specimens  with  greenish  tinge;  with  a  pair  of  widely 
separated,  narrow  brown  lines  from  anterior  median  point  of  meso- 
notum  to  ends  of  scutellum;  also,  narrow  center  of  scutellum,  palpi, 
mesopleiu"on  and  midcoxa,  narrow  apices  of  hind  femur  and  tibia, 
fifth  tarsal  segment,  and  fore,  lateral,  and  hind  margins  of  abdominal 
tergites  3  to  7,  brown.  Wing  bare  and  hyaline,  all  veins  brownish,  a 
brown  spot  over  first  radial  cell  including  Ri  and  r-m  crossvein;  a 
second  spot  over  apex  of  second  radial  cell  and  below  halfway  across 
cell  R5;  knobs  of  halteres  brown. 

Flagellar  segments  of  antenna  in  proportion  of  25 :15: 15:15:15:15: 
17:18:28:28:30:30:60.  Palpal  segments  in  proportion  of  4:15:20: 
10:22,  third  segment  not  swollen,  pit  absent.  Mesonotum  and 
abdomen  with  coarse,  dark  bristles.  Proportions  of  segments  of  hind 
leg  as  in  table  1 ;  basitarsi  without  spines,  fifth  segment  with  a  pair  of 
long,  black  batonnets  at  basal  third;  claws  slender  and  unequal,  the 
longer  as  long  as  fifth  segment,  the  other  about  half  as  long.  Sper- 
matheca  one,  large  and  ovoid,  the  duct  not  sclerotized.  Wing  with 
macro trichiae  absent;  first  radial  cell  very  small  and  narrow,  about  a 
seventh  as  long  as  second,  latter  to  0.8  wing  length;  crossvein  r-m 
forming  an  oblique  line  with  vein  Ri ;  media  petiolate  for  0.23  its  length. 

Male:  Similar  to  the  female  but  with  the  usual  sexual  differences; 
body  more  or  less  greenish  with  black  markings  as  in  female ,  antennal 
plumes  yellow  at  bases,  black  on  distal  halves.  Ninth  sternite  a 
narrow,  transverse,  anterior  band,  posterior  membrane  spiculate; 
ninth  tergite  full  and  rounded,  with  a  pair  of  low,  rounded,  setigerous, 
apicolateral  lobes.  Basistyle  about  twice  as  long  as  broad  at  base,  ab- 
ruptly narrowed  about  midway  on  inner  side,  forming  a  low,  mesal  lobe 
connected  by  an  irregular  membrane  to  ventrolateral  margins  of  ninth 
tergite;  dististyle  about  0.7  as  long  as  basistyle,  stout,  slightly  incurved, 
with  bluntly  pointed  tip.  Aedeagus  with  an  oblique  pair  of  spatulate 
sclerites  with  bases  abruptly  bent  laterad  around  bases  of  basistyles. 
Parameres  a  pair  of  broad,  flattened,  sclerotized  plates,  bases  connected 
across  mid^^ne  by  a  highly  sclerotized  band  from  outer  anterolateral 


76  PROCEEDINGS    OP   THE    NATIONAL    MUSEUM  vol.  103 

margins  of  basistyles;  base  of  each  paramere  with  a  prominent  lateral 
spur  projecting  caudad  along  inner  margin  of  basistyle;  distal  half 
greatly  expanded,  with  a  large,  sharp,  lateral  tooth;  inner  margins 
contiguous  at  apices,  each  with  a  very  slender,  pointed  spine  abruptly 
bent  ventrad  and  cephalad. 

Type.— VSNM  8356,  9,  Virginia. 

Material  examined. — Maryland:  Shadyside,  July  6,  1925,  Aldrich, 
1  9. 

Virginia:  June  16,  Pergande,  1  9  (type). 

Illinois:  Urbana,  May  20,  1914,  Aldrich,  1  cf,  19. 

Louisiana:  Baton  Rouge,  Apr.  13,  25,  1947,  Wirth,  3  cf  cf ,  3  99. 

Mexico:  Ciudad  Monte,  Tamaulipas,  Nov.  23-Dec.  ],  1943, 
Brookman,  29  cT  cf,  35  99;  Tamazanchale,  San  Luis  Potosi,  March  11, 
1944,  Brookman,  1  9;  Tapachula,  Chiapas,  Sept.  20,  1944,  Brookman, 
1  9. 

Puerto  Rico:  Arecibo,  Apr.  5,  1932,  Faxon  and  Anderson,  1  cf , 
1  9;  San  Juan,  June  14,  1933,  Harlan,  1  9. 

Jamaica:  Bath,  St.  Thomas,  Chapin  and  Blackwelder,  4  99;  Spanish 
Town,  Feb.  2,  1937,  Chapin  and  Blackwelder,  3  99. 

Panama :  Sabanas,  Apr.  20,  1923,  Shannon,  1  9. 

Remarks. — S.  fiehrigi  Kieffer,  1917,  from  Paraguay  and  Brazil, 
resembles  S.  coquilletti  very  closely  except  for  the  abdominal  markings, 
with  a  pair  of  large  lateral  brown  spots  on  the  first  tergite  and  a  large 
median  brown  spot  on  the  second,  and  tergites  3  to  7  with  dark  markings 
on  the  disc  rather  than  on  the  margins.  In  the  male  genitalia  of 
jiehrigi  the  basistyles  bear  a  prominent  mesal  lobe,  the  dististyles  are 
slenderer  at  the  apices,  the  lateral  bars  of  the  aedeagus  are  slender 
without  capitate  tips,  and  the  parameres  are  not  so  broad  and  the 
lateral  tooth  is  not  so  prominent. 

10.  Stilobezzia  (Stilobezzia)  diversa  (Coquillett),  1901 

Figure  12,  / 

Ceratopogon  diversus  Coquillett,  Proc.  U.  S.  Nat.  Mus.,  vol.  23,  p.  607,  1907  ( 9  , 

New  Jersey). 
Ceratolophus  diversus,  Kieffer,  Genera  insectoruui,  fasc.  42,  p.  60,  1906. 
Johannseniella  diversa,  Malloch,  Bull.  Illinois  State  Lab.  Nat.  Hist.,  vol.  10,  p. 

227,  1914. 
Hariomyia  diversa,  Malloch,  Bull.  Illinois  State  Lab.  Nat.  Hist.,  vol.  10,  p.  344, 

1915. 
Stilobezzia  diversa,  Johannsen,  Ann.  Ent.  Soc.  Anier.,  vol.  36,  p.  781,  1943. 

Description. — Female:  Length  1.5  mm.,  wing  1.4  mm.  by  0.6  mm. 
Head  brown,  antennae  and  palpi  yellow;  thorax  shining  black;  abdo- 
men and  coxae  light  green;  legs  yellow,  fifth  tarsal  segment  brownish; 
wings  hyaline,  anterior  veins  yellow,  halteres  j^ellowish.  Antennae  as 
long  as  entire  body;  palpi  slender,  third  segment  not  swollen.     Wing 


HELEID    MIDGES,    GENUS    STILOBEZZIA — WIRTH  77 

with  first  radial  cell  a  fifth  as  long  as  second;  petiole  of  media  a  fourth 
as  long  as  distal  portion.  Legs  slender,  nearly  bare,  proportions  of 
segments  of  hind  leg  as  in  table  1 ;  basitarsi  unspined,  fifth  segment 
with  a  pair  of  ventral  batonnets. 

Male:  Similar  to  the  female  but  with  the  usual  sexual  differences; 
plumes  of  antennae  yellow.  Ninth  sternite  about  four  times  as  broad 
as  long,  with  a  very  shallow,  broad,  posterior  excavation,  the  posterior 
membrane  spiculate;  ninth  tergite  tapered,  much  longer  than  basi- 
styles,  with  very  short,  setigerous,  apicolateral  lobes.  Basistyles  broad 
at  bases,  attenuated  distad,  simple,  with  an  irregular,  hyaline  mem- 
brane connecting  the  inner  margins  about  two-thirds  way  to  apices; 
dististyles  about  as  long  as  basistyles,  slightly  curved  and  tapered  to  a 
slender,  pointed  tip.  Aedeagus  with  a  pair  of  slender,  sclerotized, 
oblique  rods  with  straight  apices,  the  curved  bases  continuous  with 
anterior  margins  of  basistyles.  Parameres  with  bases  very  stout  and 
conical,  and  with  greatly  narrowed  apices  recurved  ventrolaterad,  the 
bases  connected  by  a  narrow,  sclerotized  band. 

Type.— Vii'NM  5482,  9,  New  Jersey,  Riverton. 

Material  examined. — New  Jersey:  Riverton,  July  3,  Johnson,  1  9 
(type). 

Virginia:  Falls  Church,  July  4,  Aug.  6,  1950,  Wu'th,  4  cf  cf ,  3  99. 

Georgia:  Thomasville,  May  15-30,  1949,  Palmer,  1  cf ,  2  99. 

11.  Stilobezzia  (Stilobezzia)  glauca  Macfie,  1939 

Figure  12,  h 

Stilobezzia  glauca  Macfie,  Rev.  Ent.,  vol.  10,  p.  204,  1939  (cf,  Brazil). — Lane,  Rev. 
Ent.,  vol.  18,  p.  207,  1947  (  9  ). 

Diagnostic  characters. — A  rather  small,  bright,  pale  green  species 
with  whitish  antennae,  legs,  wings,  and  halteres;  mesonotum  exten- 
sively dark,  shining  brown;  abdomen  with  black  band  across  third 
tergite  and  lateral  spots  on  fifth. 

Description. — Female:  Length  1.8  mm.,  wing  1.7  mm.  by  0.6  mm. 
Head  yellowish  brown,  antennae  whitish  at  bases,  last  five  segments 
and  palpi  pale  brownish.  Flagellar  segments  in  proportion  of 
25:15:15:15:16:17:18:20:40:40:35:35:45.  Palpi  long,  segments  in  pro- 
portion of  4:7:15:8:12;  third  segment  not  swollen,  pit  absent. 

Thorax  pale  green;  mesonotum  with  median  third  of  anterior  margin 
shining  brownish  black,  forming  a  broad,  V-shaped  mark  caudad,  the 
sides  of  mesonotum  broadly  dark  to  ends  of  scutellum,  and  the  median 
area  between  the  dark  bands  yellowish.  Vestiture  consisting  of  a  few 
long,  brown  bristles;  scutellum  with  four  weak,  marginal  bristles. 
Postscutellum  and  a  faint  pleural  spot  brown.  Legs  whitish,  hind 
femur  and  tibia  with  very  faint  dark,  apical  spot.  Proportions  of 
segments  of  hind  leg  as  in  table  1 ;  basitarsus  without  spines,  fifth  seg- 


78  PROCEEDINGS    OF    THE    NATIONAL    MUSEUM  vol.  103 

ment  with  a  pair  of  long,  dark  batonnets  at  basal  fourth;  claws  very 
slender  and  unequal,  the  longer  claw  as  long  as  fifth  segment,  the  other 
a  third  as  long. 

Wings  hyaline,  anterior  veins  yellowish;  first  radial  cell  small,  a 
seventh  as  long  as  second;  petiole  of  media  a  fifth  way  to  apex  of  wing. 
Halteres  yelloMash  white. 

Abdomen  pale  green;  two  dark  spots  present  on  sides  of  first  and 
second  tergites;  third  tergite  with  broad,  black  band;  fifth  tergite 
with  a  pair  of  small,  lateral,  black  spots.  Spermatheca  one,  slightly 
oval,  the  duct  not  sclerotized. 

Male:  Similar  to  the  female  but  with  the  usual  sexual  differences ; 
antennal  plumes  bright  yellow.  Ninth  sternite  broad  and  not  strongly 
sclerotized,  with  spiculate  membrane;  ninth  tergite  rounded  with 
membranous,  setigerous,  apicolateral  lobes.  Basistyle  with  distal 
portion  slender,  base  stout,  a  large  unsclerotized  lobe  about  halfway 
along  inner  margin,  connected  on  dorsal  side  to  margin  of  ninth  tergite ; 
dististyle  about  as  long  as  basistyle,  tapered  and  curved  to  a  slender, 
bluntly  pointed  tip.  Aedeagus  with  an  oblique  pair  of  long,  straight, 
slender,  sclerotized  bars.  Parameres  with  stout  basal  knobs,  a  pair 
of  inner  apodemes  joined  mesad  by  a  membrane  from  their  anterior 
margins;  stems  stout,  slightly  curved,  with  apex  abruptly  narrowed, 
pointed,  and  bent  mesad  in  the  form  of  a  short  claw. 

Type. — In  collection  British  Museum  (Natural  History),  cf, 
Brazil,  Nova  Teutonia. 

Material  examined. — Virginia:  Falls  Church,  July  4-29,  1950, 
Wu-th,  9  cf  d',  16  99. 

South  Carolina:  Greenville,  June  20,  1940,  H.  and  M.  Townes,  1  cf . 

Louisiana:  Baton  Rouge,  May  1947,  Wirth,  2  cfcf. 

Remarks. — Very  similar  to  S.  viridis  (Coquillett),  1901,  in  the  gen- 
eral green  color  with  black-banded  abdomen,  but  readily  distinguished 
from  that  species  by  the  dark-banded  mesonotum,  unhanded  legs, 
pale  wings,  and  in  the  male  by  the  yellowish  antennal  plumes.  One 
female  from  Falls  Church  is  atypical,  with  the  abdomen  entirely  dark. 

12.  Stilobezzia  {Stiloheszia)  pallidiventris  (Mallocb),  1915 

Hartomyia  pallidiventris  Malioch,  Bull.  Illinois  State  Lab.  Nat.  Hist.,  vol.  10,  p. 

344,  1915  (9  ;  Illinois,  Indiana). 
Stilobezzia  pallidiventris,  Johannsen,  Ann.  Ent.  Soc.  Amer.,  vol.  36,  p.  781,  1943. 

Description. — Female  (from  original  description):  Length  1.0  mm. 
Shining  black;  head,  antennae,  palpi,  legs,  and  abdomen  yellow; 
abdomen  may  be  darkened;  wings  hyaline;  halteres  white.  Antennae 
as  long  as  entire  body;  legs  slender;  basitarsi  longer  than  segments 
2  to  5  combined;  fifth  segment  without  ventral  batonnets;  claws 
slender,  inner  half  as  long  as  outer.  First  radial  cell  a  fifth  as  long  as 
second. 


HELEID    MIDGES,    GENUS    STILOBEZZIA — WIRTH  79 

Type. — In  collection  Illinois  Natural  History  Survey,  9,  Illinois, 
Urbana. 

Remarks. — Very  near  S.  (S.)  diver sa  (Coquillett),  but  much  smaller, 
the  abdomen  yellowish  rather  than  green,  and  fifth  tarsal  segment  of 
female  without  ventral  batonnets. 

13.  Stilobeszia  (Stilobezzia)  pruinosa  Wirth,  1952 

Figure  12,  g 

Stilobezzia  pruinosa  Wirth,  Univ.  California  Publ.  Ent.,  vol.  9,  p.  203,  1952   (cf, 
9  ;  California). 

Description. — Female:  Length  1.0  mm.  Pruinose  brown,  with 
broad,  short,  unmarked  wings  and  a  marked,  pruinose  gray  pattern 
on  the  mesonotum;  legs  with  narrow  sub-basal  pale  rings  on  tibiae; 
halteres  white.  Proportions  of  segments  of  hind  leg  as  in  table  1; 
basitarsi  and  fifth  segment  without  ventral  spines.  Spermathecae 
two,  slightly  unequal,  slightly  ovoid,  with  a  few  minute  perforations; 
the  ducts  sclerotized  a  very  short  distance. 

Male:  Ninth  sternite  transverse;  ninth  tergite  long  and  rounded 
caudad  with  a  subapical  pair  of  low,  rounded,  setigerous  lobes.  Basi- 
styles  tapered,  the  inner  margins  with  low  hump  bearing  a  few  setose 
tubercles;  dististyles  very  stout,  with  blunt  apices.  Aedeagus  with 
a  pair  of  small,  oblique,  sclerotized  bars.  Parameres  a  pair  of  highly 
sclerotized,  heavy  plates;  basal  apodemes  winglike  and  bilobed; 
inner  margins  of  stems  nearly  straight,  slightly  expanded  on  outer 
sides  about  halfway  to  apices;  apices  abruptly  narrowed  and  bent 
ventromesad  on  about  distal  fourth.  The  apices  of  the  parameres 
are  somewhat  longer  in  the  Florida  specimens  figured  than  in  the  type 
from  California. 

Types. — USNM  59945,  holotype,  cf,  allotype,  California,  San  Luis 
Obispo  County,  Arroyo  Seco  Ranger  Station. 

Material  examined. — California:  AiToyo  Seco  Ranger  Station, 
San  Luis  Obispo  County,  July  1,  1948,  Wirth,  3  cf  cf  (type,  2  para- 
types),  1  9  (allotype);  Pollack  Pines,  Eldorado  County,  July  7,  1948, 
Wirth,  1  cf ;  Snowline  Camp,  Eldorado  County,  July  8,  1948,  Wirth, 
1  (f . 

Florida:  Gainesville,  Alachua  County,  June  2,  1950,  Bidlingmayer 
1  cf ,  1  9;  Grayton  Beach,  Walton  County,  May  10,  1949,  Butler,  1  9 

14.  Stilobezzia  {Stilobezzia")  punctipes,  new  species 

Figure  11,  d 

Diagonostic  characters. — A  large,  yellowish  species  with  brown 
thorax  and  brown-punctate  legs;  abdomen  with  narrow  segmental 
dark  bands;  wings  with  two  small  black  spots. 


80  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  i03 

Description. — Female:  Length  2.4  mm.,  wing  1.9  mm.  by  0.9  mm. 
Head  pruinose  brown;  antennae  yellow,  apices  of  segments  3  to  10 
and  aU  of  last  five  segments  brown;  palpi  black.  Flagellar  segments 
in  proportion  of  28:18:18:18:18:18:20:20:38:38:40:40:55,  last 
segment  with  very  slender  pointed  tip.  Papal  segments  in  proportion 
of  5:10:16:8:20,  third  segment  slightly  swollen  with  a  small,  round, 
subapical  pit. 

Thorax  pruinose  brown,  margins  of  mesonotum,  broad  center  of 
scatellum,  and  upper  half  of  pleiu-a  yellow.  Mesonotum  with  irregular, 
silv^ery  pruinose  patches,  a  few  long,  brown  bristles;  scutellum  with 
four  long,  marginal  bristles.  Coxae  and  trochanters  dark  brown, 
rest  of  legs  yellow;  fore  femur  with  broad  sub-basal,  midfemur  with 
narrow  basal,  and  hind  femur  with  narrow  apical  brown  bands;  all 
tibiae  narrowly  brown  at  apices;  rows  of  small,  round,  brown  puncta- 
tions  at  bases  of  hairs  on  all  femora  and  tibiae,  these  hairs  long  and 
dark  on  hind  tibia.  Proportions  of  segments  of  hind  leg  as  in  table 
1;  basitarsus  with  three  black  ventral  spines  on  forelegs,  two  on 
midlegs  and  one  at  base  on  hind  legs;  fifth  tarsal  segment  without 
batonnets;  claws  slender  and  unequal,  the  longer  claw  as  long  as 
fifth  segment,  the  other  two-thirds  as  long. 

Wing  hyaline,  anterior  veins  yellowish;  a  brown  spot  over  r-m 
crossvein  and  a  very  small  one  over  vein  R2+3;  macrotrichiae  absent. 
First  radial  cell  large,  one-fourth  as  long  as  second,  latter  to  0.8  wing 
length;  petiole  of  media  as  long  as  r-m  crossvein.     Halteres  brown. 

Abdomen  dull,  pruinose  yellowish,  tergites  with  irregular  brown 
bands  in  middle  of  segments,  pleura  dark.  Spermathecae  three,  the 
two  larger  subequal  and  oval  with  short,  sclerotized  ducts,  the  third 
about  two-thirds  as  large  and  subspherical,  without  sclerotized  duct. 

Male:  Similar  to  the  female  but  with  the  usual  sexual  differences; 
plumes  of  antennae  golden  brown.  Ninth  sternite  a  very  narrow  band, 
the  posterior  membrane  spiculate;  ninth  tergite  rounded,  with  small, 
submedian,  setigerous  lobes  at  apex.  Basistyle  simple,  twice  as  long 
as  broad;  dististyle  as  long  as  basistyle  and  rather  stout  to  apex, 
slightly  bent  on  distal  third.  Aedeagus  with  an  oblique  pair  of  heavy 
twice-sinuate  sclerites  with  apices  snoutlike  and  crossing  each  other 
mesad.  Parameres  very  slender,  the  lateral  apodemes  evenly  curved 
and  slender,  stems  nearly  straight  with  bases  slightly  enlarged, 
gradually  tapered  to  sharp,  hooked  tips  abruptly  bent  ventrocephalad 
at  two-thirds  total  length. 

Ti/pes.— USNM  60966,  holotype,  9,  Crystal  River,  Citrus  County, 
Fla.,  June  7,  1949,  Hudson,  light  trap;  allotype,  same  data  except 
date,  Oct.  6,  1949.  Paratypes:  Florida:  11  d'cf,  23  99,  same  data 
as  type  except  dates,  May-October  1949-50;  1  cT,  3  99,  Leesburg,  Lake 
County,  Aug.  7,  1949,  Braddock;  Mexico:  4  99,  Ciudad  Monte, 
Tamaulipas,  Nov.  22,  1943,  Brookman. 


HELEID    MIDGES,    GENUS    STILOBEZZIA — ^WIRTH  81 

Most  closely  related  to  the  Neotropical  species  S.  (S.)  punctulata 
Lane,  1947,  and  S.  (S.)  kiefferi  Lane,  1947,  which  it  resembles  in  its 
spotted  wings  and  punctate  legs.  Both  related  species,  however, 
have  the  abdomen  reddish,  the  former  species  differing  in  having 
three  dark  wing  spots,  the  third  over  tip  of  second  radial  cell,  while 
the  second  species  has  the  first  radial  cell  half  as  long  as  the  second, 
yellowish  coxae,  whitish  halteres,  and  the  fore  femur  and  midfemur 
lack  the  basal  dark  bands. 

15.  Stilobezzia  {Stilobezzia)  rabelloi  Lane,  1947 

Figure  11,  a 

Stilobezzia  rabelloi  Lane,  Rev.  Ent.,  vol.  18,  p.  203,  1947  (c?,  9  ;  Brazil). 

Diagnostic  characters. — A  large,  brown  species  with  irregular, 
silvery,  pruinose  markings;  wing  brownish,  infuscated  with  tlu-ee 
darker  and  one  lighter  spots  on  anterior  margin;  body  hairs  dense  and 
strong;  fifth  tarsal  segment  with  3  or  4  pairs  of  ventral  batonnets. 

Description. — Female.  Length  3.0  mm.,  wing  2.2  mm.  by  0.8  mm. 
Head  pruinose  brown,  narrow  bases  of  flagellar  segments  yellow, 
palpi  black.  Antenna  with  flagellar  segments  in  proportion  of 
22:15:15:15:18:20:24:24:40:42:50:?:?  (last  two  segments  broken, 
not  measured).  Palpal  segments  in  proportion  of  8:15:25:15:20, 
third  segment  slightly  swollen,  densely  set  with  strong  hairs,  a  small, 
round  pit  near  apex. 

Mesonotum  with  strong,  conical,  anterior  spine,  the  sides  sloping, 
flattened  only  in  prescutellar  area;  color  dark  brown  with  irregular 
silvery  pruinose  markings,  usually  three  sublateral  pairs  of  small, 
rounded,  polished  spots  on  disc;  humeral  corners  yellowish;  a  few  long, 
brown  hairs  in  row.  Scutellum  brown,  yellowish  in  middle,  with 
4  or  5  long,  brown,  marginal  bristles.  Postscutellum  and  pleura  dark, 
pollinose  brown.  Legs  including  coxae  dark  brown;  fore  femur  and 
midfemur  with  narrow  sub-basal  and  subapical  and  broad  median 
yellowish  bands;  hind  femur  with  narrow,  light  band  at  apex;  fore 
tibia  and  midtibia  with  narrow  sub-basal  and  broad  median  yellowish 
bands;  first  four  tarsal  segments  yellowish.  Proportions  of  segments 
of  hind  leg  as  in  table  1 ;  leg  hairs  strong  and  spinose,  especially  long 
on  hind  tibia;  basitarsus  with  strong  black  basal  and  apical  spines,  a 
third  spine  in  middle  on  forelegs;  fifth  segment  with  3  or  4  pairs  of 
ventral  batonnets;  claws  very  slender  and  unequal,  the  longer  claw 
as  long  as  fifth  segment,  the  other  a  third  as  long. 

Wing  brownish  hyaline,  extensively  infuscated  along  the  veins; 
three  dark  brown  spots,  first  over  r-m  crossvein,  second  over  vein 
R2+3,  and  the  third  over  distal  third  of  second  radial  cell  and  extending 
halfway  through  cell  Rj  at  same  level;  a  light  area  just  beyond  tip 


82  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  103 

of  costa  as  large  as  third  browTi  spot,  the  apex  of  the  wing  beyond 
Hghtly  infuscated.  First  radial  cell  nearly  half  as  long  as  second,  the 
latter  ending  at  about  0.7  wing  length;  petiole  of  media  about  as  long 
as  first  radial  cell.     Halteres  brown  except  at  extreme  bases  of  stems. 

Abdomen  dark  brown,  tergites  with  narrow,  apical,  segmental, 
silvery  pruinose  bands;  vestitiu-e  very  dense,  consisting  of  long,  stout, 
brown  hairs.  Spermathecae  two,  ovoid,  subequal,  with  ducts  sclero- 
tized  a  short  distance,  and  a  rudimentary  third  present. 

Male:  Similar  to  the  female  but  with  the  usual  sexual  differences; 
plumes  of  antennae  bro^\^l.  Ninth  sternite  about  three  times  as 
broad  as  long,  with  shallow,  mesal,  posterior  excavation,  and  the 
membrane  bare;  ninth  tergite  short  and  rounded  with  very  short, 
setigerous,  apicolateral  lobes.  Basistyles  simple,  very  long,  slender 
and  tapered  to  tips,  about  three  times  as  long  as  broad;  dististyles 
about  0.8  as  long  as  basistjdes,  slightly  incurved,  slender,  with  pointed 
apices.  Aedeagus  with  an  oblique  pair  of  double-sinuate  sclerites 
with  mesal  tips  snoutlike.  Parameres  strongl}^  sclerotized  with 
broad,  winglike,  lateral  apodemes  at  bases,  a  lateral  pair  of  L-shaped 
or  boomerang-shaped  lateral  plates  with  blunt  apices  directed  latero- 
caudad,  and  a  submedian  pair  of  nearly  straight  processes  of  same 
thickness,  but  about  twice  as  long,  with  pointed  apices. 

Type. — -In  collection  University  of  Sao  Paulo,  Brazil,  cf,  Brazil, 
Rio  de  Janeiro,  Estrada  Rio-Sao  Paulo,  km.  47. 

Material  examined. — Georgia:  Thomasville,  ]May  15-30,  1949, 
Palmer,  1  9. 

Florida:  Leesburg,  Lake  County,  Aug,  7,  1949,  Braddock,  2  99; 
Panama  City  Beach,  May  6,  1949,  McElvey,  3  cf  cf ,  1  9;  Santa  Rosa 
Island,  Escambia  County,  June  7,  1949,  Butler,  1  cf ,  1  9. 

Louisiana:  New  Orleans,  May  27,  1947,  Van  Beeck,  6  cf  cf . 

Honduras:  La  Ceiba,  Aug.  6,  1916,  Dyer,  1  cf . 

Surinam:  Moengo,  Nov.  10,  1947,  Stage,  1  9. 

Remarks. — Closely  related  to  S.  (S.)  beckae,  new  species,  which  it 
resembles  closely  in  the  bro\vn  mesonotum  with  silvery  pruinosity  and 
strong  anterior  spinose  tubercle  and  brownish  infuscated  and  spotted 
wings,  but  beckae  differs  in  lacking  the  third  dark  spot  at  apex  of  first 
radial  cell  and  the  ventral  batonnets  on  fifth  tarsal  segment,  and  the 
body  hairs  are  not  nearly  so  well  developed. 

16.  Stilobezzia  {Stilobezzia)  sybleae,  new  species 

Figure  12,  c 

Diagnostic  characters. — A  small,  shining,  brown  species  with  dusky, 
yellow  legs;  wing  hyaline  with  a  prominent  beadlike  swelling  of  vein 
^i+5  near  apex  of  second  radial  cell;  hind  tarsi  with  fifth  segment  and 
claws  greatly  reduced. 


HELEID    MIDGES,    GENUS    STILOBEZZIA — WIRTH  83 

Description. — Female:  Length  1.5  mm.,  wing  1.3  mm.  by  0.5  mm. 
Head  shining  brown,  clypeus  and  antenna!  pedicel  yellowish,  antennal 
flagellum  and  palpi  dusky.  Thorax  polished  brown,  mesonotum  with 
sparse  black  hah's  in  rows ;  scutellum  with  four  marginal  hairs.  Pleura, 
coxae,  midfemur,  hind  femur,  and  tibiae  dusky,  yellowish  brown;  all 
of  forelegs,  midtarsus,  and  hind  tarsus  lighter  yellowish.  Proportions 
of  segments  of  hind  leg  as  in  table  1 ;  basi tarsi  unspined;  fifth  segment 
with  a  pair  of  black  ventral  batonnets  near  base;  claws  slender  and 
unequal,  normal  on  forelegs  and  midlegs  with  outer  claw  as  long  as 
fifth  segment,  the  inner  a  third  as  long,  on  hind  legs  with  fifth  segment 
half  as  long  and  claws  only  a  fourth  as  long  as  on  forelegs  and  midlegs. 

Wings  grayish  hyaline,  without  macro trichiae,  anterior  veins  brown- 
ish yellow.  First  radial  cell  a  fourth  as  long  as  second,  latter  to  five- 
eighths  wing  length ;  vein  R4+5  on  distal  half  of  second  radial  ceU  with 
an  oval  beadlike  swelling;  cross  vein  r-m  short,  petiole  of  media  about 
three  times  as  long  as  crossvein.  Halteres  brown.  Abdomen  shining 
brown,  very  convex  dorsally  and  somewhat  petiolate  at  base. 

Male:  Ninth  sternite  a  slender  anterior  band,  the  posterior  mem- 
brane spiculate;  ninth  tergite  rounded,  the  apicolateral  lobes  reduced. 
Basistyle  broad,  with  a  large  triangular,  flattened,  sclerotized  lobe 
about  midway  of  mesal  margin;  dististyle  nearly  as  long  as  basistyle, 
incurved,  gradually  tapered  to  slender  apex.  Aedeagus  with  a  pair  of 
slightly  oblique,  long,  very  slender,  sclerotized  bars.  Parameres  with 
basal  knoblike  apodemes  fused  with  anteromesal  margins  of  inner 
lobes  of  basistyles,  with  slender,  long,  curved,  sicklelike  posterior 
blades. 

Types.— USNM  60967,  holotype,  9,  and  allotype,  Falls  Church, 
Virginia,  July  4,  1950,  Wirth. 

Remarks. — This  species  is  very  similar  to  *S'.  (S.)  antennalis  (Coquil- 
lett),  but  the  highly  modified  posterior  branch  of  the  radius  and  hind 
fifth  tarsal  segment  and  claws  are  quite  distinctive.  The  West 
African  species  S.  limnophila  Ingram  and  Macfie,  1922,  is  very  similar 
in  coloration  and  in  the  structure  of  the  male  genitalia,  but  since  the 
female  is  unloiown,  the  wing  and  tarsal  features  cannot  be  compared 
with  those  of  sybleae.  Parabezzia  poikiloptera  Ingram  and  Macfie, 
1922,  from  West  Africa,  which  is  a  Stilobezzia  of  the  subgenus  Eukrai- 
ohelea,  has  a  remarkable  thickening  of  the  radius  almost  exactly  like 
that  of  sybleae,  indicating  that  this  character  is  not  just  an  aberration. 
This  species  is  named  for  my  wife,  Syble  Austin  Wirth. 

17.  Stilobezzia  {Stilobezzia)  thomsenae,  new  species 

Figure  12,  a 

Diagnostic  characters. — Very  closely  related  to,  and  almost  indis- 
tinguishable from,  bulla  Thomsen,  except  for  the  very  characteristic 
male  genitaha. 


84  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  103 

Description. — Female:  Length  1.0  mm.;  wing  0.8  mm.  by  0.4  mm. 
Apparently  somewhat  darker  brown  than  bulla,  the  mesonotum  and 
scutellum  concolorous;  femora  brownish.     Otherwise  as  in  bulla. 

Male:  Genitalia  broader  than  long,  parts  very  broad  and  stout. 
Ninth  sternite  a  narrow,  anterior  band,  posterior  membrane  spiculate; 
ninth  tergite  rounded,  with  a  submedian  pair  of  fingerlike  setose  lobes 
about  twice  as  long  as  broad.  Basistyles  simple,  about  twice  as  long  as 
broad,  set  far  apart,  no  inner  lobes;  dististyles  short  and  stout,  two- 
thirds  as  long  as  basistyle,  with  abruptly  narrowed,  pointed,  clawlike 
apices.  Aedeagus  with  anterior  arms  vestigial,  the  dorsal  median 
sclerite  a  crescent-shaped,  transverse  bar  connecting  distoventral 
angles  of  basistyles.  Parameres  stout,  with  broad,  lateral,  winglike, 
basal  apodemes,  stems  shghtly  knobbed  at  bases,  stout  and  slightly 
out-curved  toward  apices,  which  are  slightly  flattened,  expanded, 
rounded,  and  bent  ventrad. 

Type.—VSNM  609G8,  holotype,  cf,  Everglades  City,  Collier 
County,  Fla.,  Feb.  7,  1950,  Davidson,  light  trap. 

Remarks. — One  female  from  Crystal  River,  Citrus  County,  Fla., 
Sept.  18,  1950,  Hudson,  is  provisionally  referred  to  S.  (S.)  thomsenae, 
since  it  resembles  the  male  in  coloration,  although  it  may  prove  to  be 
a  dark  specimen  of  S.  (S.)  bulla.  This  species  is  named  in  honor  of 
Dr.  Lillian  Thomsen,  who  has  contributed  much  to  the  study  of 
North  American  Heleidae. 

18.  Stilobezzia  {Stilobezzia)  viridis  (Coquillett),  1901 

Figure  12,  d 
Ceratopogon  viridis  Coquillett,  Proc.  U.  S.  Nat.  Mus.,  vol.  23,  p.  607,  1901  ( 9  , 

New  Jersey). 
Ceratolophus  viridis,  Kieffer,  Genera  insectorum,  fasc.  42,  p.  61,  1906. 
Johannseniella  viridis,  Malloch,  Bull.  Illinois   State  Lab.  Nat.  Hist.,  vol.  10,  p. 

227,  1914. 
Hartomyia  viridis,  Malloch,  Bull.  Illinois  State  Lab.  Nat.  Hist.,  vol.  10,  p.  342, 

1915. 
Stilobezzia  viridis,  Johannsen,  Ann.  Ent.  Soc.  Amer.,  vol.  36,  p.  781,  1943. 

Description. — Female:  Length  2.0  mm.,  wing  1.9  mm.  by  0.6  mm. 
A  bright,  pale-green  species;  antennae  and  legs  yellow,  apex  of  antenna 
dark;  distal  fourth  of  hind  femur  and  narrow  apex  of  hind  tibia  black; 
abdomen  with  a  black  band  across  distal  half  of  third  tergite  and  a 
pair  of  large  black  spots  on  fifth  tergite.  Wings  brownish  hyaline, 
anterior  veins  infuscated.  Proportions  of  segments  of  hind  leg  as  in 
table  1 ;  basitarsi  without  spines ;  fifth  segment  with  a  pair  of  long, 
black  batonnets  at  basal  fom-th;  claw^s  slender  and  very  unequal,  the 
longer  as  long  as  fifth  segment,  the  other  about  half  as  long. 

Male:  As  in  the  female  but  with  the  usual  sexual  differences, 


HELEID    MIDGES,    GENUS    STILOBEZZIA — ^WIRTH  85 

plumes  of  antennae  dark.  Genitalia  very  large  and  bulbous.  Ninth 
sternite  reduced,  the  posterior  membrane  spiculate;  ninth  tergite 
large,  convex,  apex  rounded  with  low,  rounded,  setigerous,  apicolateral 
lobes.  Basistyle  short  and  very  broad,  nearly  as  broad  as  long; 
dististyle  about  1.2  times  as  long  as  basistyle,  extreme  base  bulbous, 
distal  portion  very  slender,  irregularly  bent  inward,  with  blunt  apex 
slightly  bent  outward.  Aedeagus  with  a  pair  of  slender,  nearly 
straight,  barlike  sclerites.  Parameres  with  reduced  basal  scleroti- 
zation,  stems  slender  and  slightly  crooked,  apices  greatly  expanded  in 
a  rounded  lobe. 

Type.—USNM  5483,  9,  New  Jersey,  Riverton. 

Material  examined. — New  Jersey:  Riverton,  June  16,  Johnson 
(type). 

Maryland:  Cabin  John,  May  26,  1943,  Cortes  and  Townes,  2  cf  cf ; 
Glen  Echo,  June  30,  1929,  Bridwell,  1  cf . 

Florida:  Leesburg,  Lake  County,  Aug.  7,  1949,  Braddock,  1  9. 

Texas:  Kirbyville,  May  20,  1908,  Tucker,  Icf. 

Remarks. — S.  glauca  Macfie  from  Brazil  and  the  southeastern 
United  States  is  closely  related  to  viridis,  but  has  the  antennae  and 
wings  infuscated,  the  mesonotum  dark,  and  lacks  the  apical  black 
markings  on  the  hind  femur. 

References 

GOETGHEBUER,   M. 

1934.  Heleidae    (Ceratopogonidae).     In    Lindner,    Die    Fliegen    der    Palal- 
arktischen  Region,  Lief.  78,  pp.  49-94. 

JOHANNSEN,  0.  A. 

1934.  New  species  of  North  American  Ceratopogonidae  and  Chironomidae. 

Journ.  New  York  Ent.  Soc,  vol.  42,  pp.  343-353. 
1943a.  Two   new  species   of   American   Ceratopogonidae    (Diptera).     Ann. 

Ent.  Soc.  Amer.,  vol.  36,  pp.  761-762. 
1943b.  A  generic  synopsis  of  the  Ceratopogonidae  (Heleidae)  of  the  Americas, 

a  bibliography,  and  a  list  of  the  North  American  species.     Ann. 

Ent.  Soc.  Amer.,  vol.  36,  pp.  763-791. 

1946.  Some  new  species  of  Nemocerous  Diptera  from  Guam.     Bull.  Bishop 

Mus.,  No.  189,  pp.  189-193. 
Lane,  J. 

1947.  Especies  Brasileiras  de  Slilobezzia  (Dipt.,  Ceratopogonidae)  e  Zygone- 

ura  stonei  nov.  nom.  (Dipt.,  Mycetophilidae) .     Rev.  Ent.,  vol.  18, 
pp.  197-214,  9  figs. 
Macfie,  J.  W.  S. 

1935.  Ceratopogonidae   (Dipt.)   from  the  river  Amazon.     Stylops,   vol.   4, 

pp.  49-56,  2  figs. 
Meillon,  B.  de 

1938.     Notes    on    African    Ceratopogonidae    (Diptera).     Proc.    Ent.    Soc. 
London,  ser.  B,  vol  7,  pp.  266-270. 

U.  S.  eoVEKNMENT  PRINTINS  OFriCEi  IMS 


PROCEEDINGS   OF   THE   UNITED   STATES   NATIONAL   MUSEUM 


SMITHSONIAN  INSTITUTION 

U.  S.  NATIONAL  MUSEUM 


Vol.    103  Washington:   1953  No.  3317 

BEETLES  OF  THE  OEDEMERID  GENUS  VASACES 
CHAMPION 


By  Ross  H.  Arnett,  Jr.^ 


Champion  (1889)  described  the  genus  Vasaces  for  three  species  from 
Central  America.  One  species,  Vasaces  aeneipe'nnis ,  was  described 
from  material  from  Mexico,  and  the  other  two,  Vasaces  sordidus  and 
Vasaces  costatus,  were  described  from  Guatemala. 

Since  the  completion  of  my  revision  of  the  Nearctic  Oedemeridae 
(Amer.  Midi.  Nat.,  vol.  45,  pp.  257-391,  1951),  I  have  seen  several 
specimens  of  Vasaces  collected  by  J.  N.  Knull  in  Arizona  and  Texas. 
This  genus  has  not  previously  been  recorded  from  North  America, 
north  of  Mexico.  In  addition,  a  few  specimens  from  Central  America, 
in  the  collections  of  the  U.  S.  National  Museum  and  the  Museum  of 
Comparative  Zoology,  have  been  available  to  me  for  study. 

The  present  paper,  based  on  all  available  material,  with  the  excep- 
tion of  the  Champion  type  specimens,  adds  four  new  species  to  the 
genus.  A  key  to  the  species  is  included,  and  the  position  of  the  genus 
in  relation  to  other  oedemerid  genera  is  discussed. 

I  wish  to  thank  Mr.  J.  N.  Knull,  Ohio  State  University,  and  Dr. 
P.  J.  Darlington,  Museum  of  Comparative  Zoology,  for  the  loan  of 
material  used  in  this  study. 

'Bureau  of  Entomology  and  Plant  Quarantine,  U.  S.  Department  of  Agriculture. 

232714—53  87 


88  PROCEEDINGS    OF    THE    NATIONAL    MUSEUM  vol.  los 

Genus  Fasaces  Champion,  1889 

Vasaces  Champion,  Biologia  Central!- Americana,  Coleoptera,  vol.  4,  pt.  2.,  p.  Ill, 
1889  [no  included  species]  and  p.  128,  pi.  6,  figs.  7  and  7,  a-c,  1890  [first  in- 
cluded species]. — Arnett,  Journ.  Washington  Acad.  Sci.,  vol.  40,  p.  225,  1950. 
(Genotype,  Vasaces  aeneipennis  Champion,  1890;  designated  by  Arnett, 
1950.) 

This  genus  belongs  in  the  subfamily  Oedemerinae  as  treated  in  my 
paper  on  the  Nearctic  species.  The  apical  segment  of  the  maxillary 
palpus  is  triangular  or  cultriform  and  the  fore  tibia  has  two  apical 
spurs.  It  is  tentatively  placed  in  the  tribe  Asclerini,  as  I  have  pre- 
viously defined  it.  However,  studies  of  certain  other  genera  have 
led  me  to  revise  somewhat  my  views  on  the  oedemerid  tribes,  and 
when  more  groups  have  been  restudied  some  rearrangement  may  be 
necessary.  Vasaces  Champion  will  key  to  couplet  7  of  my  key  to  the 
genera  of  Asclerini.  The  slightly  emarginate  eyes  separate  it  from 
Heliocis  Arnett,  and  the  triangular  or  cultriform  apical  segment  of  the 
maxillary  palpus  will  separate  it  from  the  genus  Sisenes  Champion. 
In  Sisenes  Champion  the  apical  segment  of  the  maxiUary  palpus  is 
small,  hardly  triangular.  In  addition,  many  Sisenes  species  resemble 
lycids  and  have  the  antennal  segments  somewhat  flattened,  a  condi- 
tion which  is  not  found  in  Vasaces.  In  the  key  to  the  male  genitalia, 
this  genus  runs  to  couplet  3,  where  it  may  be  separated  from  the 
genera  Eumecomera  Arnett  and  Sisenes  on  the  basis  of  the  structure 
of  the  ninth  abdominal  sternite.  In  Vasaces  the  apical  portion  of  the 
ninth  sternite  of  both  Eumecomera  and  Sisenes  consists  of  two  separate 
rods  united  only  by  thin  membrane.  The  ninth  sternite  of  Vasaces 
resembles  somewhat  that  of  Heliocis  except  that  the  base  is  two 
distinct  rods  instead  of  the  Y-shaped  piece  found  in  Heliocis. 

It  appears  that  the  genus  Sisenes  is  a  close  relative  of  Vasaces, 
both  on  characters  of  external  morphology  and  those  of  the  male 
genitalia.  However,  in  the  studied  forms,  the  two  genera  can  be 
adequately  separated  using  the  characters  mentioned  above.  In 
gross  appearance,  the  two  genera  are  very  distinct.  The  species  of 
Vasaces  are  for  the  most  part  dark,  somewhat  metallic,  elongate 
beetles,  quite  different  from  the  brilliantly  colored,  lycidlike  species 
of  Sisenes. 

Generic  diagnosis. — Head  moderate  or  elongate,  usually  constricted 
behind  the  eyes.  Eyes  somewhat  emarginate,  large.  Antenna 
slender,  11-segmented,  inserted  between  the  eye  and  the  base  of  the 
mandible,  distinctly  separated  from  the  margin  of  the  eye  and  the 
base  of  the  mandible.  Second  segment  of  the  antenna  large,  over 
one-half  the  length  of  the  fourth  and  sometimes  nearly  the  length  of 


OEDEMERID    BEETLES — ^ARNETT  89 

the  third.  Both  mandibles  with  the  apex  of  each  bifid.  Apical 
segment  of  the  maxillary  palpus  large,  triangular  or  cultriform,  widest 
at  the  base  or  near  the  center.  Thorax  slightly  longer  than  broad, 
usually  only  slightly  expanded  apically  and  only  slightly  constricted 
at  the  base.  Legs  slender,  claws  simple;  all  tarsi  with  only  the 
penultimate  segment  of  each  spongy  beneath.  Elytra  distinctly 
costate,  sometimes  markedly  so. 

Male  genitalia:  Median  lobe  short,  base  wider,  without  proc- 
esses, tapering  towards  apex,  without  apical  enlargements.  Paramere 
a  troughlike  plate,  wide  basally,  without  basal  processes,  narrowed 
toward  apex;  apex  V-shaped,  emarginate;  tegminite  large,  spatulate. 
Ninth  abdominal  sternite  hood-shaped,  with  two  basal  tails;  apex 
expanded,  with  basally  projecting  lateral  flanges.  Ninth  tergite 
two  parallel  rods,  apically  expanded  and  united  by  membrane. 
Eighth  abdominal  sternite  and  tergite  each  bilobed,  the  sternite  larger. 
Setae  present  on  the  apical  lobes  of  the  eighth  abdominal  segment. 

Biology. — Unknown. 

Distribution. — KJno^vn  only  from  Central  America  and  the  extreme 
southwestern  part  of  the  United  States. 

Key  to  the  species  of  Vasaces 

1.  Apical  segment  of  the  maxillary  palpus  cultriform   (fig.  13, gr),  its  outer  edge 

sinuate 2 

Apical  segment  of  the  maxillary  palpus  triangular  (fig.  13, /i) 4 

2.  Costae,  suture,  and  margins  of  elytra  fusco-piceous,  intervals  pale;  antennae 

uniformly    dark   except   for   small   pale   areas   at    the   joints;    legs    dark, 

slightly  paler  at  joints linearis,   new  species 

Elytra  with  uniform  coloration 3 

3.  Elytra  uniformly  brassy costatus  Champion 

Elytra  dark knulli,  new  species 

4.  Pale  testaceous;   elytra  with  costae  and  suture  each  with  an  indistinct,  inter- 

rupted row  of  fine,  oblong  streaks  on  either  side sordidus  Champion 

Dark  species,  otherwise  marked 5 

5.  Pronotum  orange  with  black  maculations,  head  short__maculatus,  new  species 
Pronotum  dark,  head  elongate 6 

6.  Pronotum  with  sides  nearly  parallel elongatus,  new  species 

Pronotum  distinctly  sinuate  laterally aenipennis,  Champion 

Vasaces  linearis,  new  species 
FiGUBB  13,  c,  e-g,  j~l,  m,  n 

Diagnostic  characters. — The  light  brassy  color,  with  the  dark  brown, 
prominent  elytral  costae,  readily  differentiates  this  species  from  the 
others.  It  is  most  closely  related  to  V.  costatus  since  the  shape  of 
the  apical  segment  of  the  maxillary  palpus  is  similar. 


90  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  i03 

Description. — (Based  on  male  holotype.)  Antenna  long,  nearly  as 
long  as  the  body,  second  segment  subequal  to  the  third.  Apical 
segment  of  the  maxillary  palpus  large,  distinctly  cultriform,  outer 
margin  arcuate.  Vertex  of  head  transversely  elevated  between  the 
eyes,  and  with  a  slight  median  groove  from  the  base  of  the  antenna 
to  the  summit  of  the  vertex.  Pronotum  somewhat  longer  than  broad, 
widest  near  anterior  margin,  slightly  narrower  at  base,  surface  irregu- 
lar, laterally  with  slight  elevations,  median  line  somewhat  impressed, 
and  with  a  slight  V-shaped  elevation  on  anterior  half.  Elytron  long 
and  narrow,  with  four  prominent  costae  and  the  suture  and  margin 
prominently  elevated,  and  a  short,  basal,  spurious  costa  uniting  with 
the  inner  margin. 

Entire  insect  clothed  with  fine,  short,  sparse,  whitish  hairs.  Color: 
in  general  pale  brassy;  head  with  a  darker  area  on  front ;  mouthparts, 
except  for  maxillary  palpi,  paler.  Antennae  and  maxillary  palpi  dark. 
Thorax  dark  except  for  irregular  paler  areas  near  base  and  apex. 
Elytra  uniformly  pale  brassy  except  for  dark  brown  costae,  suture, 
and  margins.  Ventral  surface  and  legs  dark.  Surface  irregularly 
rugose-punctate.     Length:  8  mm. 

Male  genitalia:  Described  in  the  generic  diagnosis.  The  seventh 
sternite  has  a  slight  angular  emargination  which  is  probably  a  second- 
ary sexual  character. 

Type. — Holotype,  male,  Ohio  State  Univ.  coll.,  Arizona,  Huachuca 
Mountains,  Cochise  County,  June  9,  1935,  J.  N.  Knull. 

Distribution. — Known  only  from  this  single  male  type  specimen 
from  Arizona. 

Vasaces  costatus  Champion,  1890 

Vasaces  costatus  Champion,   Biologia    Centrali-Americana,    Coleoptera,   vol.   4, 
pt.  2,  p.  129,  pi.  6,  fig.  8,  1890. 

This  species  is  also  known  only  from  one  male,  the  type,  from 
Guatemala,  at  Cerro  Zunil  (4,000  feet),  collected  by  Champion,  and 
is  in  the  British  Museum  (Natm-al  History).  It  is  much  smaller 
than  the  genotype,  but  a  little  larger  (9.5  mm.)  than  the  preceding 
species.  The  elytra  are  uniformly  brassy  in  color,  lacking  the  dark 
brown  costal,  sutural,  and  marginal  markings.  The  seventh  sternite 
of  the  male  is  deeply  emarginate. 

Vasaces  knulli,  new  species 

Figure  13,  b 

Diagnostic  characters. — The  small  size,  dark  color,  and  lack  of 
brassy  tints  serve  to  separate  this  species  from  the  other  two  having  the 
apical  segment  of  the  maxillary  palpus  cultriform.  It  is  most  closely 
allied  to  V.  costatus. 


OEDEMERID  BEETLES — ^ARNETT 


91 


Figure  13. — Adults  and  male  genitalia  of  the  genus  V  as  aces:  a,  Vasaces  maculatus,  new- 
species,  9  ;b,  F.  knulli,  new  species,  9  ;  c,  F.  linearis,  new  species,  cf ;  d,  V.  elongatus,  new 
species,  ?  ,  ^, /,  cf  genitalia,  V.  linearis,  new  species:  e,  paramere;/,  median  lobe;  g,  h, 
maxillary  palpi:  g,  V.  linearis,  new  species,  c^;  h,  V.  elongatus,  new  species,  9  ;  ^,  V- 
aeneipennis  Champion,  9  ;  j-n,  V.  linearis,  new  species,  cf  ;y,  ninth  tergite;  k,  ninth  ster- 
nite;  /,  tegminite;  m,  eighth  tergite;  n,  eighth  sternite. 


92  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  io3 

Description. — (Based  on  female  holotype.)  Antennae  short,  not 
extending  over  two-thirds  the  length  of  the  elytra,  second  and  third 
segments  equal  in  length,  each  about  three-fourths  the  length  of  the 
fourth  segment.  Apical  segment  of  the  maxillary  palpus  large, 
cultriform,  outer  edge  slightly  arcuate.  Head  shorter  than  that  of  the 
other  species;  front  with  a  slight  depression  near  the  bases  of  the 
antennae  on  the  inner  side;  a  moderately  deeply  impressed  median 
line  on  the  front.  Pronotum  with  surface  very  irregular  and  with  a 
moderately  prominent  V-shaped  ridge  on  the  anterior  half  and 
an  interrupted  lateral  ridge  on  each  side;  longer  than  broad,  widest 
anteriorly  just  beyond  the  middle,  constricted  behind.  Elytra  nar- 
row, long;  costae  present  but  obscure.  Surface  covered  with  fine, 
short,  sparse,  whitish  hairs.  Color:  dark,  mouthparts,  underside 
of  head,  neck,  coxae,  bases  of  femora,  tarsi  at  joints,  and  small  area  at 
joints  of  the  segments  of  the  antennae  paler.  Surface  finely  rugose- 
punctate.     Length:  7  mm. 

Type. — Holotype,  female,  Ohio  State  Univ.  coll.,  Arizona,  Huachuca 
Mountains,  Cochise  County,  June  9,  1935,  J.  N.  Knull. 

Distribution. — This  species  is  known  only  from  the  type,  which  is 
from  Arizona. 

Vasaces  sordiduSf  Champion,  1890 

Vasaces  sordidus  Champion,  Biologia  Centrali- Americana,  Coleoptera,  vol.  4,  pt. 
2,  p.  129,  1890. 

This  species  is  known  from  one  female  only,  the  type,  from  CapetUlo, 
Guatemala,  collected  also  by  Champion,  which  is  in  the  British  Mu- 
seum (Natural  History).  It  is  smaller  than  the  preceding  species 
(8.5  mm.) ;  the  apical  segment  of  the  maxUlary  palpus  is  smaller,  and 
triangular  instead  of  cultriform.  The  color  is  pale  testaceous.  The 
costae  and  suture  are  conspicuous,  and  each  has  an  indistinct,  inter- 
rupted row  of  fine  oblong  streaks  on  either  side. 

Vasaces  maculatus,  new  species 

FiGUBE  13,  a 

Diagnostic  characters. — The  dark  orange  pronotum  with  a  median  and 
two  lateral  spots  is  unique  among  the  species  of  this  genus.  It  is  the 
largest  known  species  in  the  genus  and  is  most  closely  related  to  V. 
sordidus  but  has  the  orange  markings  which  are  lacking  in  that  species. 

Description. — (Based  on  female  holotype.)  Antennae  short,  reach- 
ing to  about  two-thirds  the  length  of  the  elytra,  second  segment  short, 
about  two-thirds  as  long  as  the  third.  Apical  segment  of  the  maxUlary 
palpus  large,  elongate  triangular,  widest  near  basal  third,  outer  mar- 
gin straight.  Front  of  head  irregular,  with  a  circular  depression 
extending  from  the  inner  side  of  the  base  of  the  antenna  basad  about 
half  the  distance   to   the  pronotum.     Pronotum   quadrate,   widest 


OEDEMERID    BEETLES — ARNETT  93 

anteiiorly,  only  slightly  constricted  posteriorly,  siu'face  irregular  but 
without  prominent  ridges  or  depressions  except  for  a  slight  depression 
on  either  side  near  the  anterior  lateral  angle.  Elytra  long,  narrower 
apically,  about  twice  as  broad  as  thorax;  costae  present,  flat,  not  promi- 
nent. 

Entire  insect  clothed  with  very  fine,  short,  sparse,  golden  hairs. 
Surface  finely  rugose-punctate.  Head  with  punctures  larger,  more 
distinct,  with  a  stouter  recumbent  hair  arising  near  each.  Color: 
Head  orange,  with  a  large  dark  spot  on  front;  antennae  and  palpi  dark; 
thorax  orange  except  for  a  large  central  dark  spot  which  connects  with 
two  large  lateral  spots ;  legs  with  coxae  and  bases  of  femora  pale,  re- 
mainder dark ;  abdomen  and  elytra  uniformly  dark.  Length :  1 0  mm . ; 
of  paratype,  11  mm. 

Types. — Holotype,  female,  USNM  [61112,  Arizona,  Chiricahua 
Mountains,  June  8.  Hubbard  and  Schwarz.  Paratype,  female,  same 
data  as  type  except  date,  June  11. 

The  paratype  agrees  with  the  type  except  that  the  black  markings 
of  the  thorax  are  less  extensive. 

Distribution. — Known  only  from  the  two  type  specimens  from 
Arizona. 

Vasaces  elongatits,  new  species 

Figure  13,  d,  h 

Diagnostic  characters. — The  uniform  dark  color,  elongate  head  and 
thorax,  and  parallel  sides  of  the  pronotum  are  characteristic  of  this 
species.  It  is  very  different  in  general  habitus  from  the  other  members 
of  this  genus,  but  is  probably  most  closely  related  to  V.  aeneipennis. 

Description. — Head  elongate,  over  twice  as  long  as  wide  at  bases 
of  antennae,  hardly  constricted  behind  the  eyes.  Antennae  reaching 
beyond  apical  third  of  elytra,  apical  segment  of  the  maxillary  palpus 
triangular,  widest  at  middle,  outer  edge  straight.  Front  of  head 
smooth,  without  elevations  or  depressions.  Pronotum  longer  than 
broad  (7 :4) ,  surface  irregular  but  without  distinct  ridges  or  depressions, 
sides  only  slightly  sinuate.  Elytra  broad,  twice  the  width  of  the 
thorax,  tapering  behind.     Legs  long  and  slender,  costae  distinct,  flat. 

Entire  insect  clothed  with  short,  fine,  white  hairs,  those  of  the  head 
and  thorax  sparser,  but  longer  and  coarser  than  those  of  the  rest  of 
the  body.  Surface  of  head  smooth  and  shiny  with  large  distinct 
punctures  which  are  sparser  on  the  vertex;  thorax  coarsely  rugose; 
remainder  of  body  finely  rugose.  Color:  dark  fuscous;  neck,  under 
surface  of  head,  mouthparts,  and  coxae  paler.     Length:  8  mm. 

Type. — Holotype,  female,  Ohio  State  Univ.  coll.,  Texas,  Chisos 
Mountains,  Brewster  County,  June  9,  1939,  D.  J.  and  J.  N.  KnuU. 

Distribution. — KJaown  only  from  the  female  type. 


94  PROCEEDINGS    OF    THE    NATIONAL    MUSEUM  vol.  io3 

Vasaces  aeneipennis  Champion 

Figure  13,  i 

Vasaces  aenipenms  Champion,  Biologia   Centrali-Americana,  Coleoptera,  vol.  4, 
pt.  2,  p.  128,  1890. 

This  is  apparently  the  least  rare  of  the  species  in  this  genus  and 
is  one  of  the  largest  of  the  known  species.  The  triangular  apical 
segment  of  the  maxillary  palpus,  the  elongate  head,  and  the  uniform 
brassy  color  distinguish  it  from  the  other  species. 

Champion  described  this  species  from  two  specimens  (a  male  and 
a  female)  from  Totosinapan,  Mexico,  in  the  Salle  collection  (location 
unknown).  The  U.  S.  National  Museum  collection  contains  two  fe- 
males from  Guatemala,  and  the  Museum  of  Comparative  Zoology  has 
one  female  from  Honduras.  A  detailed  description  of  one  female 
from  Guatemala  follo^vs: 

Description. — (Based  on  female,  from  Guatemala).  Head  elongate, 
about  twice  as  long  as  wide  at  bases  of  antennae.  Antennae  long, 
reaching  nearly  to  apical  third  of  the  elytra,  second  segment  about 
two-thirds  the  length  of  the  third.  Apical  segment  of  the  maxillary 
palpus  triangular,  widest  at  middle,  outer  edge  straight.  Dorsal 
surface  of  head  hregular,  but  without  distinct  ridges  or  depressions. 
Pronotum  nearly  quadrate,  widest  anteriorly,  sides  distinctly  sinuate, 
surface  irregular.  Elytra  long  and  narrow,  tapering  at  apex;  costae 
distinct,  flat,  both  edges  of  costae  outlined  by  a  very  fine,  irregular, 
impressed  line,  somewhat  darker  in  color,  giving  in  gross  appearance 
the  impression  that  the  costae  are  double. 

Surface  clothed  with  extremely  fine,  very  short,  sparse,  golden 
hairs  which  are  somewhat  larger  on  the  thorax  and  dorsal  surface  of 
the  head.  Surface  rugose,  that  of  the  thorax  and  head  indistinctly 
rugose-punctate.  Color:  uniformly  brassy,  dorsal  surface  of  the  head 
and  ventral  surface  of  the  thorax  and  abdomen  darker.  Length: 
15  mm. 

Variation :  The  surface  of  the  head  and  thorax  varies  from  rugose- 
punctate  to  shiny  smooth  with  sparse  punctures.  The  length  varies 
from  10  to  16  mm.  The  color  is  darker  in  some  specimens  than  in 
others. 

Distribution. — Known  only  from  the  types  from  Mexico  and  the 
above-mentioned  specimens  from  Guatemala  and  Honduras  (Rosario 
San  Juancito). 

Material  examined.— Female,  USNM  coll.,  Guatemala,  Santa 
Maria,  W.  Schaus. 


U.   S,   GOVERNMENT  PRINTING   OFFICE:  1953 


PROCEEDINGS   OF  THE  UNITED  STATES    NATIONAL  MUSEUM 


issued  ira»?fvA. sJ^Ml  ^y  '^fi 


SMITHSONIAN  INSTITUTION 
U.  S.  NATIONAL  MUSEUM 


Vol.103  Washington  :  1953  No.  3318 

SCARABAEID  BEETLES  OF  THE  GENUS  BRADYCINETULUS 
AND  CLOSELY  RELATED  GENERA  IN  THE  UNITED  STATES 


By  O.  L.  Cartwright 


The  group  of  beetles  reviewed  in  this  paper  have  long  been  known 
under  the  name  Bolboceras.  The  species  found  in  the  United  States 
were  last  revised  by  Charles  Schaeffer,  1906  (Trans.  Amer.  Ent.  Soc, 
vol.  32,  pp.  249-254).  At  that  time  five  species  were  placed  in  Horn's 
genus  Bradycinetus,  later  incorrectly  synonymized  under  Bolboceras 
(see  synonymy  under  Bradycinetulus) ,  and  three  were  assigned  to 
Bolboceras.  Of  these  three,  Scarabaeus  lazarus  Fabricius  had  been 
placed  in  Eucanthus  by  Westwood,  1848  (Proc.  Linn.  Soc.  London, 
vol.  1,  p.  387),  and  Bradycinetus  hornii  Rivers  and  B.  minor  Linell 
were  later  removed  to  Kolbeus  by  Chapin  (see  synonymy  under 
Bolbelasmus) . 

With  the  exception  of  the  species  Eucanthus  and  Bolbocerosoma,  here 
considered  correctly  placed  in  recent  usage,  all  of  the  United  States 
species  listed  by  Schaeffer  have  been  reexamined  in  this  study  and 
reassigned  to  genera  as  required  by  the  literature  on  the  group.  Also 
included  in  this  paper  are  the  one  species  described  siiice  Schaeffer's 
time,  Bolboceras  angulus  Robinson;  a  Mexican  species  discovered  in  the 
United  States;  another  which  may  eventually  be  found  inside  our 
borders;  and  four  new  United  States  species.  Two  new  genera  are 
proposed.  A  key,  based  in  part  on  that  by  Boucomont,  is  presented 
for  the  genera  of  the  Bolboceratini  found  in  the  United  States.  The 
genera  Kolbeus,  Bolbelasmus,  Bradycinetulus,  and  the  two  new  genera 
Bolbocerastes  and  Bolborhombus  are  discussed,  synonymy  listed,  and  a 
key  to  the  species  given.  Pertinent  references,  the  type  locality  and 
location  of  the  type,  and  notes  on  distribution  follow  for  each  species. 
Distribution  is  also  shown  on  the  maps,  figures  15  and  16.     The  genus 

232990—53 1  95 


96  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  los 

Bolbocerosoma,  revised  by  Dawson  and  McColloch,  1924  (Canadian 
Ent.,  vol.  56,  pp.  9-15),  is  not  included,  nor  is  the  true  Bolboceras, 
which  was  revised  under  the  name  Odontaeus  by  J.  B.  Wallis,  1928 
(Canadian  Ent.,  vol.  60,  pp.  119-128,  151-156,  168-176). 

The  number  of  specimens  available  for  study  was  greatly  increased  by 
material  borrowed  from  museum  and  private  collections.  I  am  grate- 
ful for  the  opportunity  to  study  these  added  specimens  and  thank  the 
following  who  loaned  museum  specimens:  R.  H.  Beamer,  Snow 
Museum,  University  of  Kansas  (KAN);  E,  C.  Becker,  Canadian 
Department  of  Agriculture  (CD A) ;  M.  A.  Cazier,  American  Museum 
of  Natural  History  (AMNH) ;  P.  J.  Darlington,  Museum  of  Compara- 
tive Zoology  (MCZ) ;  Henry  Dietrich,  Cornell  University  (CU) ;  T.  H. 
Hubbell,  University  of  Michigan  (MICH);  P.  D.  Hurd,  University 
of  California  (CAL) ;  J.  N.  Knull,  Ohio  State  University  (OSU) ;  Hugh 
B.  Leech,  California  Academy  of  Sciences  (CAS) ;  L.  M.  Martin,  Los 
Angeles  County  Museum  (LAC) ;  H.  J.  Reinhard,  A.  &  M.  College  of 
Texas  (TEX);  V.  D.  Roth,  Oregon  State  CoUege  (ORE);  M.  W. 
Sanderson,  Illinois  Natural  History  Survey  (INHS) ;  and  G.  E. 
Wallace,  Carnegie  Museum,  Pittsburgh  (CARN).  My  thanks  are 
extended  also  to  those  who  loaned  specimens  from  their  private  collec- 
tions: R.  H.  Arnett,  D.  K.  Duncan,  C.  A.  Frost,  P.  C.  Grassman, 
Henry  Howden,  A.  T.  McClay,  G.  H.  Nelson,  F.  H.  Parker,  Mark 
Robinson,  E.  C.  VanDyke,  and  F.  G.  Werner.  I  am  especially  grate- 
ful to  Henry  Howden  who  examined  my  types,  compared  specimens 
with  them  and  later  furnished  those  records  marked  with  an  asterisk. 
The  few  records  of  the  eastern  Brady cinetulus  ferrugineus  (Palisot  de 
Beauvois)  taken  from  the  literature  are  marked  with  a  double  asterisk. 
Florida  records  from  the  literature  are  from  Blatchley  (Florida  Ent., 
vol.  12,  p.  29,  1925),  Alabama  records  from  Loding  (Geol.  Surv. 
Alabama  Monogr,  11,  p.  101,  1945),  and  North  Carolina  records  from 
Brimley  (The  insects  of  North  Carolina,  p.  201,  1938). 

AU  other  distribution  records  were  copied  from  labels  attached 
to  specimens  I  examined. 

Key  to  the  genera  of  Bolboceratini 

1 .  Eye  entirely  divided  by  ocular  canthus 2 

Eye  not  completely  divided  by  ocular  canthus 3 

2.  Middle  coxae  narrowly  separated;  the  intercoxal  process  with  a  vertical  tooth- 

like elevation  in  front  of  the  coxae;  species  usually  bicolored. -Bolbocerosoma 

The  intercoxal  process  without  toothlike  elevation;   species  not  bicolored; 

males  usually  with  long  cephalic  horn Bolboceras  * 

3.  Middle  coxae  subcontiguous ;  metasternum  linear  between  the  coxae 4 

Middle  coxae  separated  by  the  anterior  lobe  of  the  metasternal  plate,  which  is 

never  linear  between  the  coxae 6 

>  Odontaeus  is  a  synonym.    See  Introduction  and  also  discussion  under  Bradycinetulus. 


BOLBOCERATIlSn    OF   THE    UNITED    STATES — CARTWRIGHT         97 

4.  Elytra  with  5  discal  striae Eucanthus 

Elytra  with  7  discal  striae Bolbelasmus 

5.  Base  of  elytia  margined,  metasternal  plate  pyriform  in  outline,  pronotum 

without  postapical  carina,  first  elytral  stria  interrupted  by  scutellum 6 

Base  of  elytra  not  margined,  metasternal  plate  rhomboid  in  outline,  pronotum 
with  postapical  carina,  first  two  elytral  striae  interrupted  by  scutellum. 

Bolborhombus,  new  genus 

6.  Apex  of  tibia  of  middle  and  hind  legs  deeply  emarginate  on  outer  side,  the 

angle  adjacent  to  spurs  appearing  almost  as  a  fixed  spur;  without  prosternal 

spine Bradycinetulus 

Apex  of  tibia  of  middle  and  hind  legs  obliquely  truncate;  prosternal  spine 
behind  anterior  coxae  transverse,  doubly  pointed,  and  remote  from  acutely 
angled  intercoxal  piece Bolbocerastes,  new  genus 

Genus  Bolbelasmus  Boucotnont,  1911 

Bolbelasmus  Boucomont,  Ann.  Soc.  Ent.  France,  vol.  79  (1910),  p.  335,  1911. 
Kolbeus  Boucomont,  Ann.  Soc.  Ent.  France,  vol.  79  (1910),  p.  335,  1911. — Chapin, 
Proc.  Biol.  Soc.  Washington,  vol.  59,  p.  79,  1946. 

Chapin,  1946,  placed  Bradycinetus  hornii  Rivers  and  B.  minor 
Linell  in  the  genus  Kolbeus,  basing  his  action  on  their  close  relation- 
ship to  Kolbeus  arcuatus  (Bates).  Examination  of  specimens  of  these 
species  in  comparison  with  Bolbelasmus  gallicus  (Miilsant)  and  B. 
unicorne  (Shrank),  and  in  the  light  of  Boucomont's  key  to  the  genera, 
places  B.  minor  Linell  and  B.  hornii  Rivers  in  Bolbelasmus  rather 
than  in  Kolbeus. 

The  genus  Kolbeus  was  erected  without  a  genotype  by  Boucomont, 
1911,  for  the  species  Bolboceras  arcuatus  Bates  and  B.  coreanus  Kolbe. 
Lucas,  1920  (Catalogus  .  .  .  Coleopterorum,  p.  356),  selected 
B.  coreanus  Kolbe,  a  species  found  in  Korea  and  China,  as  the 
genotype.  The  species  B.  arcuatus  Bates  almost  exactly  duplicates 
Bolbelasmus  minor  (Linell)  except  for  the  slightly  narrower  scutellum 
and  lack  of  a  marginal  line  over  the  middle  half  of  the  base  of  the 
pronotum.  Since  a  direct  comparison  of  specimens  shows  such  minor 
differences,  I  transfer  arcuatus  Bates  to  the  genus  Bolbelasmus. 
Whether  the  genus  Kolbeus  should  be  synonymized  with  Bolbelasmus 
will  depend  upon  an  examination  of  its  genotype,  a  species  not  avail- 
able for  study  at  present.  Further  basis  for  placing  arcuatus  in  Bol- 
belasmus is  found  in  similarity  of  the  aedeagi  (see  fig.  14). 

Genotype. — Scarabaeus  gallicus  Mulsant,  Histoire  naturelle  des 
Coleopteres  de  France,  Lamellicornes,  p.  350,  1842.  (Present  desig- 
nation.) 

Key  to  the  species  of  Bolbelasmus 

1.  Base  of  pronotum  not  margined  over  middle  half;  length  7  to  9  mm.  (Cali- 
fornia,    Mexico) arcuatus    (Bates) 

Base  of  pronotum  completely  margined ,-.,. 2 


98  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.103 

2.  Ely  tral  striae  coarsely,  deeply  punctate  as  in  Eucanthus;  genae  obtusely  roundfed ; 
surface  of  head  and  clypeus  similarly  densely,  coarsely  punctate  in  male; 
frontal  carina  weakly  tri-tuberculate  in  female;  length  7  to  10  mm.  (Texas). 

minor  (Linell) 

Ely  tral    striae   finely    to   moderately   punctate;    genae   angulate   externally; 

surface  of  clypeus  roughly,  densely,  coarsely  punctate,  remaining  surface  of 

head  much  smoother  and  more  finely  punctate  in  the  male;  frontal  carina 

not  trinodose  in  female;  length  14  mm.  (California) hornii  (Rivers) 

Bolbelasmus  arcuatus  (Bates),  new  combination 

Figure  14,  I 

Bolboceras  arcuatus  Bates,  Biologia  Centrali-Amerlcana,  Coleoptera,  vol.  2,  pt.  2, 

p.  Ill,  1887. 
Kolbeus  arcuatus  Boucomont,  Ann.  Soc.  Ent.  France,  vol.  79  (1910),  p.  336, 1911. — 

Chapin,  Proc.  Biol.  Soc.  Washington,  vol.  59,  p.  79,  1946. 

Bolbelasmus  arcuatus  (Bates)  is  a  small,  dark  red-brown,  shining 
species,  7  to  9  mm.  in  length,  readily  recognized  through  the  lack  of 
basal  margins  on  both  pronotum  and  elytra.  The  labrum  is  wddely, 
shallowly  emarginate  in  front;  the  clypeus  is  almost  evenly  arcuate 
from  side  to  side;  the  clypeal  suture  is  distmct;  the  anterior  angles  of 
the  pronotum  are  sharply  right  angled;  the  scutellum  is  elongate; 
the  strial  punctures  are  coarse,  in  the  outside  row  at  the  apex  very  coarse 
and  unusually  deep,  practically  perforating  the  elytra;  the  elytra 
have  a  row  of  widely  spaced,  long,  bristlelike  hairs  near  the  margin; 
the  male  has  a  single  median  conical  cephalic  horn  and  a  low,  wide, 
median  thoracic  prominence  and  anterior  shallow  depression  on  each 
side;  the  female  has  a  wide,  more-or-less  tri-tuberculate  carina  on  the 
head  and  a  simple,  fine,  sharp,  transverse  carina  on  the  pronotum. 

Type. — Type  probably  among  the  Biologia  specimens  in  the  British 
Museum  (Natural  History). 

Type  locality. — Not  designated  among  the  localities  originally  listed, 
"Mexico,  Cordova,  Toxpan,  Playa  Vicente  (Salle),  Colima  City 
(Hoge);  Nicaragua,  Chontales  (Belt,  Janson)." 

Specimens  examined. — 19, 

Distribution. — Costa  Rica:  Tilaran  (Guanacaste),  San  Pedro  de 
Montes  de  Oca,  San  Jose,  and  *Dominical.  Guatemala.  Mexico: 
Colima,  El  Sabino,  Compostela,  MazatMn,  Monterey,  *Monclova, 
and  *Guadalajara.  United  States:  a  single  specimen  labeled  "Los 
Angeles,  California,  H.  Klages  Coll.,"  in  the  Carnegie  Museum, 
Pittsburgh,  Pa. 

Figure  14. — Ventral  and  lateral  views  of  the  aedeagi  of  Bradycinetulus  and  closely  related 
species:  a-c,  Species  of  Bolborhombus:  a,  parvulus,  new  species;  b,  angulus  (Robinson); 
c,  schaefferi  (Boucomont).  d-g,  Species  of  Bolbocerastes:  d,  peninsularis  (Schaeffer);  e, 
regalis,  new  species;!  f,  serratus  (LeConte);  g,  imperialis,  new  species,  h-j.  Species  of 
Bradycinetulus:  h,  ferrugineus  (Palisot  de  Beauvois);  i,  rex,  new  species;  j,  fossatus 
(Haldeman).  k-n,  Species  of  Bolbelasmus:  k,  gallicus  (Mulsant);  /,  arcuatus  (Bates); 
m,  hornii  (Rivers);  n,  minor  (Linell). 


BOLBOCERATINI    OF    THE    UNITED    STATES — CARTWRIGHT         99 


L     ^  V y  M  fg 

Figure  14. — (For  explanation  see  facing  page.) 


100  PROCEEDINGS    OF    THE    NATIONAL   AIUSEUM  vol.  los 

Bolbelasmus  minor  (Linell),  new  combination 

Figure  14,  n 

Bradycinetus  minor  Linell,  Proc.  U.  S.  Nat.  Mus.,  vol.  18,  p.  723,  1895. 
Bolboceras  minor  Schaeffer,  Trans.  Amer.  Ent.  Soc,  vol.  32,  p.  253,  1906. 
Kolbeus  minor  Chapin,  Proc.  Biol.  Soc.  Washington,  vol.  59,  p.  79,  1946. 

Bolbelasmus  minor  (Linell)  almost  exactly  duplicates  B.  arcuatus 
(Bates)  except  that  it  averages  slightly  larger,  the  punctures  are 
coarser  throughout,  and  the  pronotum  is  completely  and  distinctly 
margined  basally.  It  is  a  small,  dark  red-bro-wn,  shming  species, 
7  to  10  mm.  in  length;  the  labrum  is  widely,  shallowly  emarginate  in 
front;  the  clypeus  is  almost  evenly  arcuate  from  side  to  side;  the 
clypeal  suture  is  distinct;  the  anterior  angles  of  the  pronotum  are 
sharply  right  angled;  the  scutellum  is  elongate;  the  strial  punctures 
are  very  coarse,  those  at  the  apex  of  the  elytra  being  tubular  and 
so  deep  as  to  nearly  perforate  the  elytra ;  near  the  margin  of  the  elytra 
is  a  row  of  widely  spaced,  long,  bristlelike  hairs;  the  male  has  the 
head  and  clypeus  similarly  densely  punctate,  the  head  bears  a  median, 
conical  horn,  and  the  pronotum  has  a  low,  wide,  median  prominence 
with  anterior,  shallow  depressions  each  side;  the  female  has  a  wide, 
transverse,  tri-tuberculate  carina  on  the  head  and  a  simple,  fine, 
transverse  caruia  on  the  pronotum. 

Type.— Rolotyipe  female,  USNM  562. 

Type  locality. — San  Diego,  Tex. 

Specimens  examined. — 2 1 . 

Distribution. — Texas:  Hidalgo  County,  Weslaco,  Brownsville,  Kings- 
ville,  and  San  Diego. 

Season. — May  16  to  June  16. 

BolbelasTnus  hornii  (Rivers),  new  combination 

Figure  14,  m 

Bradycinetus  hornii  Rivers,  Bull.  California  Acad.  Sci.,  vol.  2,  p.  61,  1886. 
Bolboceras  hornii  Schaeffer,  Trans.  Amer.  Ent.  Soc,  vol.  32,  p.  253,  1906. 
Bradycineiulus  hornii  Cockerell,  Ent.  News,  vol.  17,  p.  242,  1906. 
Kolbeus  hornii  Chapin,  Proc.  Biol.  Soc.  Washington,  vol.  59,  p.  79,  1946. 

Bolbelasmus  hornii  (Rivers)  is  quite  similar  to  B.  minor  (Linell)  but 
is  considerably  larger,  lighter  in  color,  and  has  much  finer  strial 
punctures.  The  scutellum  is  not  so  elongate,  being  nearly  an  equi- 
lateral triangle  with  arcuate  sides.  Other  characters  given  in  the 
key  should  readily  identify  this  species. 

Type. — Location  of  type  unknown  to  me.  (According  to  Dr. 
VanDyke  the  entire  Rivers  Collection  was  purchased  by  Dr.  Walther 
Horn  and  it  is  presumably  in  the  Berlin-Dahlem  collection.) 

Type  locality. — "Sonora,  Tuolumne  Co.,  Cal.  Found  also  in 
Sacramento  Co." 


BOLBOCERATINI    OF    THE    UNITED    STATES — CARTWRIGHT      101 

Specimens  examined. — 349. 

Distribution. — California:  Sequoia  National  Park,  Napa,  Kaweah, 
Camp  Greely  (Tulare  County),  Mendocino,  Mokelumne  Hill,  Camp 
Greely  (Fresno  County),  Three  Eiders,  El  Dorado  County,  Coulter- 
v^ille,  Sonoma  County,  Potwisha,  San  Joaquin  Experiment  Station 
(Madera  County),  Mount  Diablo,  Sacramento,  Mineralking,  La  Jon 
Pass,  Kattlesnake  (Placer  County),  *Cala7eras  County,  and  *0'Neals 
(Madera  County).    Baja  California:  Hamilton  Ranch. 

Season. — January  19  to  May  8,  with  greatest  number  in  April. 

Genus  Brady cinetulus  Cockerell,  1906 

Bolboceras  Kirby  (in  part),  Trans.  Linn.  Soc.  London,  vol.  12,  pt.  2,  p.  459,  1818. 

(Type  not  designated.) 
Amechanus  Horn  (not  Thomson,  1864),  Trans.  Amer.  Ent.  Soc,  vol.  3,    p.  48, 

1870. 
Bradycinetus  Horn  (not  Sars,  1865),  Trans.  Amer.  Ent.  Soc,  vol.  3,  p.  334,  1871. 
Bradycellus  Schaeffer  (not  Erichson,  1837),  Trans.  Amer.  Ent.  Soc,  vol.  32,  p. 

249,  1906. 
Bradycinetulus  Cockerell,  Ent.  News,  vol.  17,  p.  242,  1906.    (Type  not  designated.) 

The  genus  Bolboceras  was  erected  by  Kirby  in  1818  for  the  eight 
species  Scarabaeus  mobilicornis  Linnaeus,  S.  mobilicornis  vax.  tes- 
taceus  Fabricius,  S.  guadridens  Fabricius,  S.  farctus  Fabricius,  S. 
lazarus  Fabricius,  S.  cyclops  Olivier,  *S'.  cephus  Fabricius,  and  Bol- 
boceras australasiae  Kirby.  Technically  no  genotype  was  designated 
although  he  wrote,  "My  details  of  Bolboceras  were  taken  from  B. 
quadridens."  Nearly  all  of  the  species  mentioned  have  since  been 
moved  to  other  genera.  Curtis,  1829  (British  Ent.,  vol.  1,  pt.  1,  p. 
74),  selected  the  species  Scarabaeus  mobilicornis  Fabricius  as  the  type 
of  Bolboceras.  Therefore,  since  our  species  of  Odontaeus  are  con- 
generic with  mobilicornis,  they  now  take  the  generic  name  Bolboceras 
and  it  becomes  necessary  to  find  an  available  name  for  those  species 
we  have  formerly  placed  in  Bolboceras.  A  search  of  the  literature 
indicates  that  name  must  be  Bradycinetulus,  as  shown  in  the 
synonymy. 

Characters  for  Bradycinetulus,  in  addition  to  those  listed  in  the  key 
to  genera,  include  the  following:  The  mandibles  are  evenly  arcuate  ex- 
ternally; the  median  prominence  of  the  pronotum  is  developed  into  a 
pair  of  horns  in  the  male;  the  scutellum  is  wider  than  long  and  dis- 
tinctly punctate  with  fine  to  moderate  punctures;  and  the  posterior 
vertical  face  of  the  prosternal  intercoxal  piece  is  wide  and  flat,  with 
the  ventral  edge  evenly  arcuate  or  slightly  angulate  at  middle.  The 
three  species  placed  here  are  all  large,  17  to  21  mm.  in  length  by  10 
to  12  mm.  in  width.  They  are  sufficiently  differentiated  in  the  follow- 
ing key. 

Genotype . — Scarabaeus  ferrugineus  Palisot  de  Beauvois,  1 809 .  (Pres- 
ent designation.) 


102  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  los 

Key  to  the  species  of  Bradycinetulus 

1.  Eye  canthus  nearly  straight  edged  laterally,  wide  clypeal  horn  of  male  bisinuate 

apically,  the  external  angles  projecting  forward,  acute,  and  scarcely  thicker 
than  the  adjacent  edges,  which  converge  posteriorly;  median  thoracic  horns 
heavy,  directed  upward,  their  sharply  rounded  tips  recurved  posteriorly 
(North    Carolina   to    Florida  and    Mississippi). 

ferrugiiieus  (Palisot  de  Beauvois) 
Eye  canthus  notched  or  emarginate,  not  straight  edged  laterally 2 

2.  Wide  clypeal  horn  of  male  distinctly  trinodose  apically,  the  external  angles 

not  projecting  bej^ond  median  point,  lateral  edges  parallel;  median  thoracic 
horns  heavy,  projecting  forward  and  outward,  sharply  rounded  and  very 

little  recurved  (Nebraska  to  Texas  and  Arkansas) fossaius  (Haldeman) 

Apex  of  clypeal  horn  of  male  widely  truncate,  straight  across,  the  external 
angles  slightly  nodose,  lateral  edges  parallel  over  apical  two-fifths,  then 
diverging  to  base;  median  thoracic  horns  slender,  directed  forward  and  up- 
ward, basally  flattened  in  front  and  behind   (Texas) rex,  new  species. 

Bradycinetulus  ferrugineus  (Palisot  de  Beauvois),  1809 

Figure  14,  h:  Plate  3 

Scarabaeus  ferrugineus  Palisot  de  Beauvois,  Insectes  recueillis  en  Afrique  et  en 

Amerique,  livr.  6,  p.  90,  1809. 
Bolboceras  lecontei  Dejean,  Catalogue  des  Coleoptfeies  .  .  .,  ed.  3,  vol.  3,  p.  149, 

1833. 
Athyreus  ferrugineus  Klug,  Abh.  Berlin  Acad.,  1843,  p.  22. 
Bolboceras  ferrugineus  Lacordaire,  Histoire  naturelle  des  insectes,  vol.  3,  p.  143, 

1856. 
Amechanus  ferrugineus  Horn,  Trans.  Amer.  Ent.  Soc,  vol.  3,  p.  48,  1870. 
Bradycinetus  ferrugineus  Horn,  Ent.  Amer.,  vol.  1,  p.  89,  1885. 
Athyreus  (Bradicinetus)  ferrugineus  Boucomont,  in  Wytsmanu,  Genera  insectorum, 

fasc.  7,  p.  8,  1902. 
Bradycinetus  ferrugineus  Schaeffer,  Trans.  Amer.  Ent.  Soc,  vol.  32,  p.  250,  1906. 
Bradycinetulus  ferrugineus  Cockerell,  Ent.  News,  vol.  17,  p.  242,  1906. 
Bolboceras  (Amechamus)  ferrugineus  Boucomont,  Ann.  Soc.  Ent.  France,  vol.  79, 

p.  341,  1910. 

Type. — Location  of  type  unknown  to  me. 

Type  locality. — South  CaroHna. 

Specimens  examined. — 107. 

Distribution. — North  Carolina:  West  End,  Southern  Pines,  **Beau- 
fort,  *Hamlet,  *Carolina  Beach,  and  **Tarboro.  South  Carolina: 
Meredith,  Florence,  Windsor,  Johns  Island,  and  Bulls  Island.  Geor- 
gia: Bainbridge,  Spring  Creek  (Decatur  County),  and  Augusta.  Flor- 
ida: Port  Saint  Joe,  Kissimmi,  Orlando,  Sanford,  Enterprise,  Miami, 
Crescent  City,  Indian  River,  Cedar  Keys,  *Lutz,  *Wacissa,  *Gaines- 
ville,  **Saint  Augustine,  **LaGrange,  and  **Dunedin.  Alabama: 
**Lee  County,  and  **Mobile  County.  Mississippi:  Ocean  Springs, 
Biloxi,  and  Leakesville. 

Season. — March  11  to  September  23,  with  the  largest  number  of 
records  in  June  and  July. 


BOLBOCERATINI    OF    THE    UNITED    STATES — CARTWRIGHT      103 

Bradycinetulus  fossatus  (Haldeman),  1853 

Figure  14,  j;  Plate  3 

Bolboceras  fossatus  Haldeman,  Proc.  Acad.  Nat.  Sci.  Philadelphia,  vol.  6,  p.  362, 

1853. 
Amechanus  fossatus  Horn,  Trans.  Amer.  Ent.  Soc,  vol.  3,  p.  48,  1870. 
Bradycinetus  fossatus  Schaeffer,  Trans.  Amer.  Ent.  Soc,  vol.  32,  p.  250,  1906. 
Bradycinetulus  fossator  Cockerell,  Ent.  News,  vol.  17,  p.  242,  1906. 
Bolboceras   {Amechamus)   fossatus   Boucomont,    Ann.  Soc.  Ent.  France,  vol.  79, 

p.  341,  1910. 

In  a  few  male  specimens  of  Bradycinetulus  fossatus  (Haldeman)  the 
outer  cariniform  edges  bordering  the  thoracic  fossae  show  a  slight 
tendencA^  to  break  near  the  upper  end  to  form  a  second  inner  carina,  a 
step  tow^ard  the  condition  found  in  Bolbocerastes. 

Type. — In  LeConte  collection,  MCZ. 

Type  locality. — Texas. 

Specimens  examined. — 102. 

Distribution. — Nebraska:  Imperial.  Kansas:  Reno  County,  Me- 
dora,  and  Sylvia.  Oldahoma:  Noble  County,  Cleveland  County, 
Payne  County,  Thomas,  Cleo  Springs,  Ferris,  and  *Alva.  Arkansas: 
Pine  Bluff.  Texas:  Fedor,  Henrietta,  Tyler,  Jacksonville,  Gold- 
thwaite,  Robinson,  Lexington,  Jefferson,  College  Station,  Lindale, 
Bexar  County,  Dallas,  Victoria,  Columbus,  Paris,  Austin,  San  Antonio, 
and  Morris  County. 

Season. — May  16  to  August,  with  most  records  in  June. 

Bradycinetulus  rex,  new  species 

Figure  14,  i;  Plate  3 

Holotype  male, length  21  mm.,  width  13  mm.  In  general  appearance 
verj^  similar  to  Bradycinetulus  jerrugineus  (Palisot  de  Beauvois)  and 
B.  fossatus  (Haldeman),  rufo-testaceous,  moderateh^  shining,  densely 
hairy  beneath.  Ch^peus  perpendicular  in  front,  the  nearly  flat,  verti- 
cal face  extendmg  upward  to  form  a  broad  truncate  horn  about  one- 
fourth  wider  than  high.  The  face  and  sides  of  the  horn  reticulate, 
the  enclosed  spaces  concave,  shallow,  and  coarse;  the  truncate  apex 
scarcely  thickened,  with  the  external  angles  very  slightly  nodose; 
the  sides  parallel  over  apical  two-fifths,  diverging  posteriorly;  margin 
thin,  irregularly  serrated,  elevated  just  before  the  eye  canthus  to  a 
very  moderate  right-angled  prominence,  the  posterior  edge  of  which 
continues  posteriorly  as  a  low,  sharp  carina  separating  the  eye  canthus 
from  the  flattened  discal  surface  of  the  head.  Canthus  sloping  away 
from  the  discal  level  of  head;  anterior  and  lateral  margins  sinuate  to 


232990—53- 


104  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  los 

the  rounded,  elevated,  anterior  angle,  surface  ridged  inward  from  the 
angle,  depressed  along  anterior  edge  and  in  posterior  angle.  Margin 
of  head  along  inner  edge  of  the  eye  with  a  reflexed  carina,  which  grad- 
ually curves  away  from  the  eye,  increases  to  a  maximum  height,  then 
angles  sharply  inward  and  backward  and  sharply  descends  to  general 
level  of  head ;  head  surface,  including  eye  canthi,  reticulate-tuberculate- 
punctate,  very  rough  anteriorly  on  the  concave  posterior  of  the  clypeal 
horn,  gradually  smoother  over  discal  area,  which  is  bounded  poste- 
riorly by  a  low,  rounded  ridge  between  the  eyes;  the  reticulation  and 
tubercles  of  the  discal  area  flatten  until  the  outlines  of  the  coarse, 
shallow  punctures  are  barely  traceable ;  head  behind  the  ridge  smooth 
at  middle  but  gradually  roughened  near  the  eyes. 

Pronotum  similar  to  that  of  B.  ferrugineus  (Palisot  de  Beau  vols) 
but  with  the  horns  of  the  median  prominence  and  the  high  lateral 
edges  of  the  depressions  much  farther  forward.  The  anterior  edge 
of  the  pronotum  very  finely  margined,  the  middle  third  arching  slightly 
over  the  base  of  the  head,  each  outer  third  without  a  curve,  straight, 
diverging  anteriorly  to  the  right-angled  anterior  angles,  slightly 
concave  close  along  these  straight  edges,  then  very  obtusely  convex 
and  rounding  into  the  deep  lateral  depressions,  the  middle  third  with 
a  smaller,  closer,  parallel  convexity  and  a  moderate  concavity  at  each 
end;  pronotum  laterally  with  two  very  large  deep  fossae  on  each  side, 
the  lower  wider  and  less  deep,  widely  sinuating  the  lateral  margin 
from  middle  to  near  the  anterior  angles,  the  upper  very  deep,  the 
outer  edge  thrown  up  in  a  high,  thin,  curving  edge  which  descends 
to  the  posterior  boundary  of  the  lower  cavity;  posterior  angles  distinct 
but  obtusely  rounded;  sides  and  base  margined,  the  edge  rather  finely 
serrate  laterally;  pronotum  convex  basally;  disc  forming  a  broad 
median  prominence  with  a  sharp  horn  pointing  upward  on  each  side, 
the  horns  being  rather  sharply  undercut  below  and  basally  flattened 
in  front  and  behind;  a  deep  median  line  from  base  forward  over  the 
median  prominence  and  down  between  the  horns.  Surface  of  the 
pronotum  alutaceous  and  finely  scabrous  back  of  the  anterior  margin, 
especially  at  the  sides,  then  shallowly,  coarsely  punctate  over  the 
inner  lower  surface  of  the  lateral  fossae;  posterior  angles  similarly 
finely  alutaceous  and  closely,  coarsely,  shallowly  punctate;  elsewhere 
finely  to  minutely,  sparsely  punctate  throughout,  except  for  the  very 
smooth  surface  of  the  deeper  fossae;  and  with  a  large  group  of  close 
very  coarse  punctures  on  each  side  back  of  the  horn  and  above  the 
deeper  fossa,  another  group  following  the  median  line  and  spreading 
anteriorly  to  bases  of  the  horns. 

scutellum  finely  punctate.    Elytra  not  strongly  shining;  minutely 


BOLBOCERATIIsTI    OF    THE    UNITED    STATES — CARTWRIGHT      105 

alutaceous  under  high  magnification;  margined  at  base;  striae  mod- 
erately impressed,  the  strial  punctures  fine,  moderately  deep;  first 
stria  interrupted  at  scutellum;  the  second  indistinct  near  scutellum. 

Metasternal  lobe  separating  the  middle  coxae  half  as  wide  as  the 
plate  behind  the  coxae;  lateral  edges  sharply  carinate. 

Female  unknown. 

Type.— Iloloiype  male,  USNM  61455. 

Type  locality. — Sarita,  Tex.  Holotj^pe  collected  on  bare  sand, 
Nov.  30,  1911. 

Paratypes. — One  male,  CAS,  Corpus  Christi,  Tex.,  June  28,  1942, 
E.  S.  Ross;  one  male,  in  Mark  Robinson  collection,  Kingsville,  Tex., 
C.  T.  Reed. 

Remarks. — Brady cinetulus  rex  may  be  readily  separated  from  the 
two  closely  allied  species  B.  ferrugineus  (Palisot  de  Beauvois)  and  B. 
fossatus  (Haldeman)  by  the  obvious  differences  in  the  clypeal  and 
thoracic  horns.  It  may  be  noted  also  that  the  last  tAvo  species  have 
only  a  single  deep  fossa  on  each  side  of  the  pronotum,  while  in  B.  rex 
this  is  doubled,  the  lower  cavity  sinuating  the  lateral  margin. 

Bolbocerastes^  new  genus 

This  genus  is  closely  allied  to  Bradycinetulus  but  differs  in  that  the 
apex  of  the  tibia  of  the  midleg  and  hind  leg  is  obliquely  truncate,  the 
scutellum  is  smooth  or  minutely  punctate,  the  mandibles  are  parallel, 
nearly  straight  sided  and  bent  sharply  inward  anteriorly,  the  sides  and 
anterior  edges  forming  a  rectangle,  the  median  prominence  of  the 
pronotum  is  without  horns,  two  elevated  carinae  are  present  on  each 
side  of  the  pronotum,  the  prosternal  spine  behind  the  anterior  coxae 
is  transverse,  doubly  pointed  and  remote  from  the  acutely  angled 
intercoxal  piece,  and  the  aedeagus  of  the  male  is  of  a  characteristic  form 
(see  fig.  14). 

Genotype. — Bolbocerastes  regalis,  new  species. 

Key  to  the  species  of  Bolbocerastes 

1.  Anterior  clypeal  horn  of  male  widely  truncate;  anterior  and  median  horns 

of  female  usually  equal  in  width,  smooth   back  of  median   horn  (Arizona, 

Nevada,  California) regalis,  new  species 

Anterior  clypeal  horn  of  male  narrowly  truncate  or  rounded;  anterior  horn 
narrower  than  median  horn  in  female,  punctate  back  of  median  horn.        2 

2.  Dull,  dark  reddish  brown;  elytral  striae  extremely  fine,  not  impressed,  the 

intervals  completely  flat  even  in  reflected  light,  strial  punctures  very  fine; 
fully  developed  male  with  the  anterior  clypeal  horn  narrowly  rounded  and 
very  little  advanced,  much  closer  and  more  nearly  in  line  with  the  strongly 
developed  lateral  horns  than  in  serratus  and   imperialis    (California,  Baja 

California) peninsularis  (SchaefFer) 

Elytral  striae  fine  to  moderate,  usually  at  least  slightly  impressed,  the  intervals 
weakly  convex  in  reflected  light;  fully  developed  males  not  as  above 3 


106  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  los 

3.  Lateral  serrations  of  pronotum  arising  between  the  two  edges  of  a  doubly 

carinate  margin;  elytral  striae  moderately  impressed,  strial  punctures  dis- 
tinct; color  reddish  brown,  usually  quite  strongly  shining;  tip  of  clypeal 
horn  rounded  in  male,  inner  thoracic  carina  without  secondary  angle  at 
upper  end;  female  cephahc  horn  rarely  higher  than  clypeal  horn,  carina 
extending  back  from  clypeal  horn  on  each  side  sharp,  strong,  usually  dark 
colored,  lateral  secondary  horns  nearer  midway  between  clypeal  and  cephalic 
horns  than  to  the  cephalic  horn,  discal  area  of  head  distinctly  punctate 
throughout,  fine  punctures  of  median  thoracic  prominence  distinct  and 
noticeable  (Kansas,  Oklahoma,  Texas,  New  Mexico). serratus  (LeConte) 
Pronotum  without  or  with  only  traces  of  a  doubly  carinate  margin  laterally, 
the  serrations  appearing  as  thickenings  of  a  single  sharp  edge;  well  developed 
males  with  a  secondary  angle  at  upper  end  of  inner  thoracic  carina —       4 

4.  Elytral  striae  much  less  distinct,  the  elytra  smoother  in  appearance,  nearly 

as  in  peninsularis ,  color  lighter,  more  yellow-brown,  less  strongly  shining; 
fully  developed  male  with  tip  of  clypeal  horn  narrowly  truncate  and  well 
forward  from  the  lateral  horns,  upper  lateral  thoracic  carina  of  the  male 
frequently  with  a  noticeable  secondary  angle  at  upper  end;  female  median 
cephalic  horn  often  very  high,  usually  higher  than  clypeal  horn,  carina 
extending  back  to  lateral  secondary  horn  weak,  sometimes  obhterated,  not 
darker  in  color,  the  lateral  secondary  horns  weak,  usually  much  farther 
back,  more  nearly  in  line  with  the  cephalic  horn,  discal  area  of  head  usually 
with  only  scattered  punctures  behind  median  horn  (Baja  California,  Cali- 
fornia, Arizona,  western  Texas) imperialis,  new  species 

Elytral  striae  deeper,  color  red-brown,  more  strongly  shining  as  in  serratus, 
otherwise  as  above  (Kansas) imperialis  kansanus,  new  subspecies 

Bolbocerastes  regalis,  new  species 

Figure  14,  e;  Plate  4 

Holotype  male,  length  18  mm.,  width  11.5  mm.  In  general  appear- 
ance closely  similar  to  Bolbocerastes  serratus  (LeConte),  ferrugineus, 
shining,  densely  hairy  beneath.  Clypeus  perpendicular  in  front,  the 
flat  vertical  face  extending  upward  to  form  a  broad  truncate  horn, 
the  vertical  height  subequal  to  base,  the  lateral  edges  arcuate  from 
base  to  upper  edge,  narrowing  at  basal  third  to  half  the  basal  width 
then  expanding  to  three-fourths,  the  upper  edge  weakly  emarginate 
and  subparallel  with  base,  shining  black  in  color, rounded  and  knoblike 
at  each  end;  the  sharply  defined  lateral  triangular  faces  of  the  horn 
concave  and  posteriorly  elevated  from  base  into  secondary  horns  half 
as  high;  posteriorly  the  broad  curving  slope  of  the  clypeal  horn  merges 
into  the  flattened  upper  surface  of  the  head,  this  quadrangular  surface 
with  a  secondary  horn  at  each  anterior  angle  and  with  sharply  defined 
carinalike  subparallel  lateral  margins  from  which  the  prominent, 
widely  explanate  eye  canthus  slopes  downward  and  outward.  Occip- 
ital area  smooth;  gradually,  finely,  closely,  shallowly  punctate 
anteriorly;  anterior  and  lateral  faces  of  clypeal  horn,  labrum,  eye 
canthi,   and  upper  surface  of  mandibles  finely  reticulate-scabrous. 


BOLBOCERATESn    OF    THE    UNITED    STATES — CARTWRIGHT      107 

Broadly  rounded  labrum  twice  as  wide  as  long.  Upper  surface  of 
mandibles  concave,  sides  straight,  bent  at  right  angles  in  front, 

Pronotum  retuse  in  front;  the  deeper  lateral  impressions  extending 
upward  on  each  side  of  the  median  prominence  and  bounded  laterally 
by  two  carinae  on  each  side;  the  lower  external  carina  starting  near 
the  serrate  lateral  thoracic  margin  and  extending  upward  parallel 
with  the  basal  thoracic  margin  for  a  distance  equal  to  half  the  length 
of  the  lateral  margin;  the  upper  carina  starting  inside  the  upper  end 
of  the  lower  and  extending  upward  for  a  slightly  shorter  distance; 
viewed  from  the  side  the  lower  carina  is  nearly  straight,  the  upper 
end  with  a  gentle  downward  curve,  the  lower  end  with  a  noticeable 
but  low  toothlike  elevation;  in  outline  the  upper  carina  is  half  as 
high  as  long,  sloping  upward  from  a  sharp  basal  curve  to  a  rounded 
right-angled  peak,  then  down  on  a  concave  curve  to  a  noticeable  sec- 
ondary angle.  The  median  pronotal  prominence  sharply  declivous  in 
front,  upper  surface  divided  by  an  anteriorly  increasingly  depressed 
median  line,  the  parallel  sides  with  sharply  rounded  edges  anteriorly; 
retuse  front  impunctate  except  for  a  subapical  line  of  close,  weak-to- 
obsolete  punctures;  median  prominence  with  fine,  close  punctures 
shading  to  minute  punctation  across  base;  a  few  obsolete  moderate 
punctures  in  the  shallow  lateral  depressions  outside  the  carinae;  basal 
bead  distinct.  Lateral  edges  strongly  serrate,  as  in  Bolbocerastes 
serratus  (LeConte). 

Elytra  not  strongly  shining;  striae  scarcely  impressed;  strial 
punctures  fine,  moderately  close,  intervals  practically  flat,  very 
minutely  punctulate. 

Metasternal  lobe  separating  middle  coxae  half  as  wide  as  plate 
behind  the  coxae. 

Female  allotype,  length  21  mm.,  width  10  mm.  Similar  to  the  male 
except  that  the  thoracic  prominences  are  much  less  developed  and 
the  head  is  different.  The  vertical  face  of  the  clypeus  is  only  half  as 
high  as  the  basal  length,  the  upper  edge  less  than  half  the  basal  length, 
and  from  each  end  a  curving  carina  extends  outward  to  top  of  the 
secondary  horn  or  tubercle;  from  the  flattened  area  back  of  and  about 
twice  as  high  as  the  secondary  tubercles  arises  a  widely  truncate 
median  horn;  anterior  to  this  median  horn  the  surface  is  reticulate- 
scabrous,  posteriorly  it  is  smooth. 

7V2)e.— Holotype  male,  USNM  61076. 

Type  locality. — The  holotype  bears  the  following  label  data:  "3206. 
Colo.  R.  bottom.  Monument  204.  Mex.  Bd.  line  Mar.  20-31,  '94. 
USNM  Ace.  No.  28133  Dr.  E.  A.  Mearns."  Monument  204  is  approx- 
imately 20  miles  south  of  Yuma,  Ariz.,  near  San  Luis,  Mexico,  its 
exact  location  being  lat.  32°  29'  03.59",  long.  114°  46'  44.79". 


108  PROCEEDINGS    OF    THE    NATIONAL    MUSEUM  vol.  io3 

Allotype  and  paratypes. — Allotype  and  97  paratypes  bear  data  as 
follows:  Nevada:  1,  Hawthorn,  June  8,  1943,  Wallace;  1,  Mount 
Charleston,  8,000-10,000  feet,  Aug.  6,  1944,  Zinn. 

California:  2,  Baker,  Mar.  27,  1935,  and  April  1935,  Saylor;  1, 
California  (State  label  only);  3,  Coachella,  May  12,  13,  17,  1927,  Van 
Duzee;  3,  Coachella,  Riverside  County,  Apr.  28,  1928,  May  9,  1927, 
July  15,  1927,  Wymore;  2,  Coachella  Valley,  May  9,  1927,  Van  Dyke, 
Aug.  20,  1949,  V.  d.  Bosch;  1,  Coyote  Wells,  Imperial  County,  June  5, 
1949,  Hurd;  1,  Essex,  Apr.  29,  1937;  1,  Holtville,  June  28,  1936, 
Cazier;  14,  Imperial  County,  Carrizo  Light,  Mar.  14,  17,  18,  22,  1928, 
Searl;  1,  Indian  Wells,  Riverside  County,  Apr.  25,  1939,  Martin; 
8,  junction  of  Highways  78  and  99,  Imperial  County,  Apr.  15,  1944, 
Tieman;  4,  Mecca,  Mar.  25,  1922,  Benedict;  3,  Murrays  Dam,  San 
Diego  County,  Cottle;  1,  Needles,  Apr.  1,  1941,  Linsley;  2,  Palm 
Springs,  May  20,  1905,  May  9,  1932;  2,  San  Clemente  Island,  Apr.  2,  3, 
1939,  Channel  Islands  Biological  Survey,  Von  Bloeker;  1,  San  Diego 
County,  Mar.  10,  1928,  light,  Well;  7,  Yermo,  San  Bernardino  County, 
Apr.  11,  1936,  Comstock,  Sept.  6-30,  1939,  Pierce. 

Arizona:  7,  Arizona  (State  label  only);  4,  Ehrenberg,  Oct.  18,  1938, 
Mar.  21,  23,  30,  1939,  Parker;  6  (includes  allotype),  Gila  Bend,  Ma-.  1, 
1930,  Lebart;  1,  Holbrook,  May  5,  1941,  Dukes;  1,  Navaho  County, 
Duncan;  1,  Phoenix;  1,  Red  Rock,  Spring  1930,  Burdett;  1,  Tucson, 
April  192-,  Duncan;  11,  Yuma,  February  1910,  Carlson,  Mar.  12,  18, 
1912,  Slevin,  May  15,  1939,  Aitken. 

Mexico,  Sonora:  1,  Choya  Bay,  Mar.  27,  1949,  Bradt;  1,  30  miles 
southwest  of  Sonoyta,  500  feet.  Mar.  31,  1949,  Bradt;  4,  Puerto 
Libertad,  Mar.  15,  1939,  Parker. 

Paratypes  in  AMNH,  CAS,  LAC,  KAN,  MCZ,  CAL,  CU,  OSJ, 
MICH,  INHS,  USNM,  and  the  private  collections  of  Mark  Robinson, 
Henry  Howden,  C.  A.  Frost,  A.  T.  McClay,  F.  H.  Parker,  D.  K. 
Duncan,  and  R.  H.  Arnett. 

Remarks. — Bolbocerastes  regalis  varies  from  16  to  20  mm.  in  length 
and  from  10  to  12.5  mm.  in  width.  It  is  closely  similar  to  B.  serratus 
(LeConte)  but  the  males  differ  noticeably  in  having  a  higher,  wide, 
truncate  clypeal  horn,  smoother,  less  punctate  pronotum,  and  on  each 
side  a  doubly  angulate  inner  pronotal  carina  which  is  always  higher 
in  the  middle.  In  B.  serratus  (LeConte)  the  anterior  clypeal  horn  is 
bluntly  rounded,  the  pronotal  punctures  are  more  evident  throughout, 
and  the  inner  lateral  pronotal  carina  is  not  doubly  angulate  or  higher 
in  the  middle.  The  truncated  tops  of  the  anterior  and  median  horns 
are  about  equal  in  width  in  females  of  B.  regalis  and  the  whole  anterior 
part  of  the  head,  including  clypeal  and  cephalic  horns,  appears  higher 
than  the  posterior  part,  like  an  elevated  snout.  The  lateral  horns  are 
most  prominent  in  this  species. 


BOLBOCERATINI    OF    THE    UNITED    STATES CARTWRIGHT      109 

Bolbocerastes  imperialis,  new  species 

Figure  14,  g;  Plate  4 

Holotype  male,  length  19  mm.,  width  11  mm.  Very  similar  to 
Bolbocerastes  serratus  (LeConte) ;  ferrugineus;  moderately  shining; 
hairy  beneath.  Vertical  face  of  clypeal  horn  inclined  forward  about 
30  degrees  from  perpendicular,  slightly  convex  when  viewed  from  side, 
height  seven-tenths  width  at  base,  width  of  the  truncate  apex  slightly 
less  than  half  the  basal  width,  the  apex  arcuate  posteriorly  at 
each  side;  the  posterior  face  of  the  horn  concave  from  side  to  side 
between  the  wealdy  carinate  edges,  which  converge  very  slightly  from 
apex  to  base.  General  level  of  the  head  slightly  more  than  half  the 
height  of  the  clypeal  horn  above  the  wide,  truncate  labrum;  this  flat- 
tened area  sloping  upward  posteriorly  and  bounded  by  a  ridge  or 
carina  deeply  arcuate  posteriorly  from  the  anterior  edge  of  one  eye  to 
the  other  and  continuing  forward  on  each  side  to  the  tips  of  the  weakly 
elevated,  right-angled,  secondary  horns,  then  converging  to  the  an- 
terior edge  of  the  clypeal  horn  and  forming  the  upper  edge  of  the  tri- 
angular lateral  area  at  base  of  the  horn;  flattened  area  of  head  slightly 
depressed  laterally  above  the  prominent  quadrangular  eye  canthus, 
which  slopes  arcuately  downward,  outward,  and  finally  upward  at  the 
outer  angles;  entire  surface  in  front  of  the  curving  posterior  ridge 
densely,  rouglily  punctate,  smooth  back  of  the  ridge.  Mandibles 
concave  dorsally;  strongly  angled  front  and  back. 

Pronotum  retuse  in  front  and  with  serrate  lateral  margins  as  in  B. 
serratus  (LeConte)  and  B.  regalis;  the  median  prominence  and  lateral 
carinae  nearly  as  in  B.  regalis  but  with  the  outer  carina  with  the  upper 
end  curving  slightly  away  and  not  quite  parallel  with  the  basal  mar- 
gin when  viewed  from  the  side;  the  inner  lateral  carina,  viewed  from 
the  side,  with  the  lower  end  rising  abruptly  to  a  right  angle,  then 
sloping  evenly  upward  to  a  right-angled  peak  at  middle,  then  down 
in  a  strongly  sinuate  in-and-out  curve;  both  carinae,  viewed  from 
directly  in  front,  evenly  arcuate,  with  the  peak  angle  of  the  inner  carina 
strongly  bent  inward;  the  median  prominence  slightly  constricted 
toward  the  base,  with  the  sides  weakly  angulate  in  front  of  the  con- 
striction and  the  surface  more  broadly  and  less  deeply  concave  from 
side  to  side  than  in  B.  regalis  and  evenly  arcuate  from  front  to  back;  a 
distinct  median  line  evident  basally;  the  apical  and  deeper  lateral 
pronotal  depressions  extending  upward  between  the  lateral  carinae; 
the  median  prominence  impunctate  except  for  half  a  dozen  or  so  close, 
moderate  punctures  on  each  side  at  base  of  the  vertical  sinus;  above 
these  about  twenty  or  so  scattered  fine  punctures  on  the  inner  slope 
of  the  vertical  sinus,  and  a  very  few  fine  punctures  indicating  a  sub- 
apical  line;  median  prominence  and  lateral  basal  area  back  of  the 


110  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  103 

carinae  closely,  finely  punctate;  smoother  and  more  finely  punctate 
across  base  at  middle;  groups  of  moderate  punctures  in  the  depressions 
back  of  the  carinae  and  an  irregular  row  of  close  moderate  punctures 
just  inside  the  basal  bead. 

Scutellum  closely,  minutely  punctate.  Elytra  one-sixth  wider  than 
long;  extremely,  finely  alutaceous  under  moderate  magnification; 
striae  scarcely  impressed;  strial  punctures  fine,  distant  about  four 
diameters;  intervals  with  very  minute  scattered  punctures. 

Metasternal  lobe  separating  middle  coxae  one-third  as  wide  as  the 
plate  behind  the  coxae. 

Allotype  female,  length  14  mm.,  width  9  mm.  Similar  to  the  male 
except  that  the  pronotal  prominences  are  less  strongly  developed  and 
the  head  is  different.  The  vertical  face  of  the  clypeus  is  one-third 
as  high  as  basal  length;  its  upper  truncate  edge  also  one-third  the 
length  of  the  base;  the  concave  triangular  sides  of  the  clypeal  horn 
with  the  upper  edge  about  twice  as  long  as  the  anterior  edge;  the 
wide  truncate  frontal  horn  is  about  three-fourths  as  high  as  wide  and 
about  four  times  as  high  as  the  low  secondary  lateral  horns,  or  tubercles. 

7V^^.— Holotype  male,  USNM  61077. 

Type  locality. — Imperial  County,  Calif.,  "on  the  Experiment  Farm," 
June  1912,  J.  C.  Bridwell. 

Allotype  and  paratypes. — Allotype  female  and  210  paratypes  bear 
the  following  data:  Texas:  1,  Castolon,  June  1,  1928,  Bibby;  1, 
Tornilla  Flat,  Big  Bond  National  Park,  July  12,  1948,  desert,  3,650 
feet,  at  light,  Werner  and  Nutting;  2,  Boquillas,  Brewster  County, 
July  7,  1948,  C.  and  P.  Vaurie;  1,  Marathon,  June  14,  1948,  D.  Rocke- 
feller Expedition,  Cazier;  1,  Reeves  County,  June  10,  1939,  Robinson. 

New  Mexico:  1,  Artesia,  July  29,  1937,  D.  J.  and  J.  N.  Knull;  1, 
White  Sands,  July  23,  1933,  Benedict. 

Arizona:  4,  Agua  Fria,  Aug.  6,  1917,  Bequaert,  June  26,  1938, 
Duncan;  2,  Aquila,  Maricopa  County,  Aug.  21,  22,  1927,  CU  lot  542, 
sub  330;  2,  Arlington,  July  25,  1948,  Anderson;  15,  Arizona  (State 
label  only);  5,  Avondale  Ranch,  Agua  Fria  River,  Aug.  7,  1917, 
Wheeler  leg.;  4,  Baboquivari  Mountains,  July  24,  1941,  Beamer, 
July  24,  1941,  Todd;  2,  Buckeye,  July  30,  31,  1935,  at  light,  Christen- 
son;  1,  Cashion,  Maricopa  County,  Aug.  25,  1949,  at  light,  Werner 
and  Nutting;  5,  Cibola,  August  1911;  1,  Continental,  Pima  County, 
July  29,  1948,  Ball;  1,  Coyote  Mountains,  Aug.  4,  7,  1916;  1,  Dome, 
July  21,  1924,  Van  Duzee;  4,  Ehrenberg,  July  20,  1946,  June  12, 
1943,  Grassman,  July  18,  1939,  Stager,  June  21,  1938,  Parker;  10, 
Globe,  Duncan,  Sept.  8,  1930,  July  11,  1931,  July,  August,  Duncan 
and  Parker;  7,  Hope,  Yuma  County,  Aug.  12, 1948,  greasewood  desert, 
1,400  feet,  at  light,  Werner  and  Nutting;  1,  Hot  Springs,  July  16 
(year  not  given),  at  light,  MorrUl;  2,  Huachuca  Mountains;  2,  Kits 


BOLBOCERATIIsTI    OF    THE    UNITED    STATES — CARTWRIGHT      111 

Peak,  Rincon,  Baboquivari  Mountains,  Aug.  1,  4,  1916,  about  1,050 
feet;  1,  Littlefield ;  1,  Marinette,  Aug.  4,  1918,  Schiffel;  1,  Olberg, 
Sept.  13,  1936,  Crandall;  1,  Palmerlee;  8,  Phinax  (sic),  September 
and  November  1921;  17,  Phoenix,  Sept.  12,  1935,  Crandell,  May  12, 
1934,  Parker,  May  29,  1922,  Odell,  Aug.  2,  1917,  CU  Biol.  Expedition, 
May  17,  1944,  Parker,  Apr.  29,  1941,  Parker,  May  7,  1918,  Bradley, 
Sept.  12,  1926;  1,  Pima  County,  Sept.  1,  1925,  Marsh;  1,  Prescott;  2, 
Sacoton,  Oilman;  1,  SafFord,  September  (no  date  given),  Duncan; 
2,  San  Bernardino  Ranch,  Cochise  County,  3,600  feet,  sycamore- 
willow  in  valley  at  light,  July  25,  1949,  Werner  and  Nutting,  July  16, 
1949,  light,  Parker;  1,  San  Luis,  Yuma  County,  Aug.  11,  1940;  1, 
Talklai,  Pollock;  20,  Tucson,  July  21  (no  year  given)  Hubbard  and 
Schwarz,  July  30,  1937,  Parker,  August  1912,  Breitenbecker,  Aug.  20, 
1932,  Flock,  August  1935,  Wharton,  October  1927,  Hamilton  (2,500 
feet),  July  10,  1950,  Aug.  8,  1950,  Bradt,  Oct.  14,  1919;  3,  Wellton, 
June  13,  1939,  at  light,  Stitt;  1,  Welton,  Aug.  9,  1917,  Wheeler  leg.; 
1,  Yuma. 

California:  7,  Blythe,  May  19,  1942,  Allen,  Aug.  26,  Hurd,  April 
16,  1945,  Constantine,  May  19,  1943,  Vargas,  July  22,  1947,  July  8, 
1947;  4,  Blythe,  Riverside  County,  July  24,  1947,  Aug.  24,  1947, 
MacSwain;  2,  California  (State  label  only);  29,  Coachella,  Riverside 
County,  Apr.  28,  1927,  May  9,  12,  17,  28,  1927,  Wymore,  May  22, 
1928,  Van  Dyke,  May  13,  1917,  Van  Duzee;  2,  Coachella  Valley, 
Sept.  8,  1929,  Comstock,  Aug.  20,  1949,  v.  d.  Bosch;  1,  El  Centro, 
Aug.  28,  1948,  light  trap;  8,  Holtville,  June  1936,  Ross,  August  (no 
date  given),  Craig,  June  28,  1936,  Cazier,  June  27,  1936,  July  15, 
1936;  16  (includes  allotype).  Imperial  County,  June  1912,  on  Experi- 
ment Farm,  Bridwell;  4,  Imperial  County,  June  1925,  Field,  July  1, 
1925,  Gehring;  2,  Indio,  July  17,  1923,  Benedict,  May  23,  1919, 
Slevin;  2,  Meloland,  June  4,  7,  1949,  v.  d.  Bosch;  1,  Needles,  June 
12,  1940,  Barr;  1,  Neighbors,  Apr.  27,  1930,  Hornung;  1,  Palo  Alto, 
July  28,  1916;  1,  Vallecito,  San  Diego  County,  Sept.  15,  1945,  at  light, 
Martin;  2,  Warners,  June  18,  1925. 

Mexico:  3,  10  miles  south  of  Catavina,  Baja  California,  July  29, 
1938,  Michelbacher  and  Ross. 

Paratypes  in  CAS,  LAC,  AMNH,  CARN,  INHS,  CAL,  OSU,  CU, 
MICH,  TEX,  KAN,  CDA,  USNM,  and  the  private  collections  of 
R.  H.  Arnett,  D.  K.  Duncan,  C.  A.  Frost,  P.  C.  Grassman,  Henry 
Howden,  A.  T.  McClay,  Gayle  H.  Nelson,  F.  H.  Parker,  Mark 
Robinson,  and  F.  Werner. 

Remarks. — Males  of  B.  imperialis  vary  from  11  to  19  mm.  in  length, 
and  from  7  to  11  mm.,  in  width.  In  lesser  developed  males  the  punc- 
tures become  fewer  and  weaker  over  the  posterior  third  inside  the 
arcuate  carina  at  back  of  the  head,  anteriorly  they  persist  strongly; 


112  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  i03 

the  clypeal  horn  decreases  in  height  and  its  apex  becomes  narrow, 
sometimes  rounded,  but  usually  evidently  truncate,  and  the  anterior 
face  vertical  or  sloping  slightly  backward;  the  median  pronotal  promi- 
nence is  usually  not  constricted  or  narrowed  basally. 

B.  imperialis  is  allied  to  B.  serratus  (LeConte),  B.  regalis  Cartwright, 
and  to  B.  peninsularis  (Schaeffer),  which  I  believe  is  a  distinct  species. 
Well-developed  males  of  B.  imperialis  are  separated  from  the  other 
species  by  the  anterior  clypeal  horn  being  relatively  farther  forward 
from  the  lateral  horns,  the  triangle  formed  by  the  tips  of  the  three 
horns  approaching  an  equilateral  triangle  much  more  closely  than  in 
the  other  species.  The  apex  of  the  clypeal  horn  is  much  narrower 
than  in  B.  regalis,  and  usually  truncate  rather  than  rounded  as  in 
serratus  and  peninsularis;  in  fully  developed  males  the  inner  lateral 
carina  is  doubly  angulate  as  in  regalis  but  has  the  high  middle  peak 
bent  inward,  while  in  specimens  of  serratus  and  peninsularis  this  carina 
is  not  higher  or  angulate  at  middle.  The  color  of  serratus  and  penin- 
sularis is  dark  red-brown,  of  imperialis,  a  more  yellow  brown. 

In  Bolbocerastes  peninsularis,  represented  by  13  specimens,  3  of 
them  cotypes,  the  male  clypeal  horn  is  rounded  as  in  B.  serratus  but  is 
much  farther  back  relative  to  the  lateral  horns,  which  are  nearly  as 
high  as  the  clypeal  horn.  The  female  is  more  heavily  punctate,  as  in 
serratus,  and  the  area  back  of  the  median  horn  is  much  more  noticeably 
punctate  than  in  imperialis. 

Bolbocerastes  imperialis  kansanus,  new  subspecies 

Holotype  male,  length  17  mm.,  width  11  mm.  Very  similar  to  B. 
imperialis;  moderately  shining ;  darker  in  color;  hairy  beneath.  Anterior 
face  of  the  narrowly  truncate  clypeal  horn  vertical ;  lateral  secondary 
horns  more  prominent;  surface  back  of  the  clypeal  horn  with  a  notice- 
able swelling  on  each  side  and  diagonally  backward  from  the  posterior 
ridges  of  the  horn  to  a  point  even  with  the  forward  edge  of  the  eye 
canthus,  leaving  between  them  a  widening  depressed  area  which 
merges  with  the  general  posterior  level  of  the  head;  surface  anterior 
to  the  arcuate  ridge  between  the  eyes  more  concave  and  much  smoother 
than  in  typical  imperialis ;  occiput  smooth;  eye  canthus  with  a  swelling 
diagonally  inward  from  outer  anterior  angle. 

Pronotum  more  coarsely  punctate  throughout;  lateral  edges  of  the 
median  prominence  more  nearly  cariniform  and  not  constricted 
basally;  median  peak  of  upper  lateral  carina  not  bent  inward:  the 
depression  back  of  the  anterior  angles  at  bottom  of  the  lateral  sinuses 
deeper,  more  noticeable,  and  distinct;  serrations  of  the  lateral  margin 
appearing  as  thickenings  of  a  single  sharp  edge;  otherwise  similar  to 
typical  imperialis. 

Elytral  striae  deeper,  nearly  as  in  B.  serratus  (LeConte) ;  the  second 
stria  basally  indistinct  over  twice  the  length  of  the  scutellum. 


BOLBOCERATINI    OF    THE    UNITED    STATES — CARTWRIGHT      113 

Aedeagus  as  in  imperialis. 

Allotype  female,  length  16  mm.,  width  11  mm.  Similar  to  typical 
B.  imperialis  except  that  punctm-es  are  coarser  throughout;  foveae 
are  deeper;  elytral  striae  are  deeper;  and  the  eye  can  thus  has  a  convex 
swelling  diagonally  inward  from  the  anterior  angle. 

Type. — Holotype  male,  in  Canadian  Department  of  Agriculture 
collection. 

Type  locality. — Kush  County,  Kan. 

Allotype  and  paratypes. — Allotype  and  6  paratypes  bear  data  as 
follows:  Kansas:  4  (including  allotype  female),  Eush  County,  Aug.  5, 
1920;  2,  Ness  County,  Aug.  5,  1920;  1,  Ness  County,  2,260  feet, 
July  5,  1912,  Williams.  Paratypes  vary  from  13  to  17  mm.  in  length, 
and  8  to  11  mm.  in  width. 

Paratypes  in  CD  A,  KAN,  and  USNM. 

Remarks. — Bolbocerastes  imperialis  kansanus  may  be  separated  from 
B.  serratus  (LeConte)  by  the  doubly  angulate  upper  lateral  thoracic 
carina  in  well-developed  males,  by  the  diagonal  convexity  of  the  eye 
canthus,  by  the  lateral  serrations  of  the  pronotum  appearing  as 
thickenings  of  a  single  sharp  margin,  and  by  the  second  elytral  stria 
all  but  disappearing  at  twice  the  length  of  the  scutellum.  In  serratus 
the  serrations  of  the  pronotal  margin  arise  between  the  two  edges  of 
the  doubly  carinate  margin  and  the  second  elytral  stria  becomes 
indistinct  opposite  the  apex  of  the  scutellum. 

Bolbocerastes  serratus  (LeConte),  new  combination 

Figure  14,  /;  Plate  4 

Alhyreus  serratus  LeConte,  Proc.  Acad.  Nat.  Sci.  Philadelphia,  vol.  7,  p.  80,  1854. 
Amechanus  serratus  Horn,  Trans.  Amer.  Ent.  Soc,  vol.  3,  p.  48,  1870. 
Bradycinetus  serratus  Horn,  Proc.  California  Acad.  Sci.,  ser.  2,  vol.  4,  p.  334,  1894. 
Athyreus  (Bradicinetus)  serratus  Boucomont,  in  Wytsmann,  Genera  insectorum, 

fasc.  7,  p.  8,  1902. 
Bradycinetus  serratus  Schaeffer,  Trans.  Amer.  Ent.  Soc,  vol.  32,  p.  251,  1906. 
Bradycinetulus  serratus  Cockerell,  Ent.  News,  vol.  17,  p.  242,  1906. 
Bolboceras  (Amechamus)  serratus  Boucomont,  Ann.  Soc.  Ent.  France,  vol.  79, 

p.  341,  1910. 

Bolbocerastes  serratus  (LeConte)  and  B.  imperialis  Cartwright,  ex- 
cept for  fully  developed  males,  are  frequently  difficult  to  separate. 
The  locality  label  will  help  in  most  cases  since  serratus  is  found  east  of 
New  Mexico,  while  imperialis  is  found  in  California,  Arizona,  and 
rarely  into  west  Texas.  The  maps  (figs.  15  and  16)  show  the  known 
distribution.  In  doubtful  cases  the  male  genitalia  will  separate  the 
two  (see  fig.  14,/  and  g).  In  serratus,  the  basal  side  or  fl^oor  of  the 
cavity  is  formed  by  the  two  sides  approaching  each  other  with  knife- 
edges,  sharply  angulate  near  the  outside  margin  (in  Bolbocerastes  the 
aedeagus  of  the  male  is  hooded  apically  by  the  two  sides  coming  together 


114 


PROCEEDINGS    OF    THE    NATIONAL    MUSEUM 


•    Bolbocerastes   imperialis  Cortwright  x    Bradyclnetulus    rex    Cartwright 

o    Bradyclnetulus    fossotus  (Holdeman) 


n^^ 

^ 

-^^ 

A^ 

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^M 

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— V^ 

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e)    1    \ 

x/~X 

>'^-'H™^ 

\  *  \     7 

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^^         T*~/S> 

L    \ 

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•    Bolborhombus    schoefferi  (Boucomont)        o    Bolbelosmus   orcuatus  (Bates) 
X     Bolbocerastes    penlnsularis   (Schoeffer) 

Figure  15. — Distribution  of  species  of  Bradyclnetulus  and  close'y  related  species. 


BOLBOCERATINI    OF    THE    UNITED    STATES — CARTWRIGHT      115 


e     Bolbelasinus  horni  (Rivers) 
M     Bolbelosmut  minor  (Linell) 


•  Bradycinetulus  ferruginous  (Palisot  de   Beouvois) 

♦  Bolbocerastes    Imperialis  Itansanus  Cartwright 


•  Bolbocerostes    regalls   Cortwrlght  o    Bolborhombus    ongulus  (Robinson) 

•  Bolbocerostes    serrotus  (Leconte)  x     Bolborhombus    porvulus   Cortwrlght 

Figure  16. — Distribution  of  species  of  Bradycinetulus'^&nd  closely  related  species. 


116  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  io3 

to  form  a  deep  cavity) ;  in  imperiaJis,  these  edges  are  rounded  and  lead 
straight  in  without  a  break  or  angle. 

Type. — In  LeConte  collection,  MCZ, 

Type  locality. — Mexican  Boundary,  Laredo  to  Ringgold  Barracks. 

Specimens  examined. — 181. 

Distribution. — Kansas:  Clark  County.  Oklahoma:  Taloga,  Chey- 
enne, *Alva,  and  *Payne  County.  Texas:  Brownsville,  *Beeville, 
Burnet  County,  Cypress  Mills,  Comal  County,  Cameron  County, 
Caterina,  Colorado  City,  Cotulla,  Childress,  Dimit  County,  Dallas, 
Dallas  County,  Edinburg,  Eagle  Pass,  Fedor,  Forestburg,  Frio  State 
Park,  Goldthwaite,  Hidalgo  County,  Kingsville,  Laredo,  Memphis, 
Menard,  *Mineral  Wells,  New  Braunsfels,  Oakville,  Plesanton,  Reeves 
County,  Round  Mountain,  Sabinal,  Taloga,  Uvalde,  Weslaco,  and 
Zavalla  County.  New  Mexico:  State  label  only  (H.  W.  Wenzel  col- 
lection). Mexico:  La  Gloria,  south  of  Monclova,  Coahuila;  and 
Rancho  Preso,  Nuevo  Le6n. 

Season. — April  28  to  September  25,  with  largest  numbers  collected 
in  May,  June,  and  September. 

Bolbocerastes  peninsularis  (Schaeflfer),  new  combination 

FiQDRE  14,  d;  Plate  4 

Bradycinetus  serratus  var.  peninsularis  Schaeffer,  Trans.  Amer.  Ent.  Soc,  vol.  32, 

p.  252,  1906. 
Bolboceras  serratus  var.  peninsularis  Boucomont,  in  Schenkling,  Coleopterorum 

catalogus,  pt.  46,  p.  13,  1912. 

All  specimens  of  this  species  examined  have  been  dull,  smooth,  and 
of  a  uniform  very  dark  red-brown  color.  The  male  is  quite  distinctive 
(see  pi.  4,  lower  right).  In  the  female  the  moderately  high  cephalic 
horn  is  twice  the  width  of  the  tip  of  the  clypeal  horn  and  the  sec- 
ondary lateral  horns  or  tubercles  are  usually  more  prominent  than  in 
B.  imperialis.  The  carinae  between  these  and  the  clypeal  horn  are 
strong  and  well  marked. 

Type. — Lectotype  male,  present  selection,  USNM  42568. 

Type  locality. — Santa  Rosa,  Baja  California,  Mexico. 

Specimens  examined. — 13. 

Distribution. — California:  2,  State  label  only,  CARN.  Mexico: 
Santa  Rosa,  San  Felipe,  La  Paz,  and  San  Ignacio,  all  in  Baja  Cali- 
fornia. 

Season. — July  27  to  October  1. 

Bolborhombus^  new  genus 

Mandibles  arcuate  externally.  Head,  male  and  female,  without 
median  frontal  horn.  Pronotum  serrate  laterally;  apex  with  a  carina 
paralleling  anterior  margin,  the  interspace  interrupted  behind  each 


I 


BOLBOCERATESri    OF    THE    UNITED    STATES — CARTWRIGHT      117 

eye  by  a  distinct  rather  deep  foveola;  base  margined.  Scutellum  tri- 
angular, with  base  straight,  sides  arcuate.  Elytra  not  margined  at 
base;  seven  discal  striae,  the  first  two  interrupted  by  the  scutellum, 
the  second  sometimes  indistinctly  forked  opposite  the  apex  of  scutel- 
lum. Two  prosternal  spines  in  tandem  behind  anterior  coxae,  the 
posterior  more  or  less  hastate.  Metasternal  plate  rhomboid  in  shape, 
the  posterior  angle  acute  and  with  adjacent  edges  cariniform,  the 
lateral  angles  with  adjacent  edges  rounded;  the  intercoxal  lobe  deeply 
concave  and  with  strong  cariniform  lateral  edges. 

Genotype.^ — Brady  cinetus     carinatus     Schaeffer,  1906  =  Bolboceras 
schaefferi  Boucomont. 

Key  to  the  species  of  Bolborhombus 

1.  Clj'peus  of  male  carinately  margined  in  front,  the  very  wide  clypeal  horn 

arcuate  anteriorly,  a  low  sharp  carina  arcuate  in  opposite  direction  behind 
the  clypeal  horn,  which  is  midway  between  this  and  the  carinate  anterior 

margin;  female  unknown angulus  (Robinson) 

Male  clypeus  not  distinctly  carinate  along  anterior  edge  but  with  parallel 
carinae  extending  back  from  anterior  angles  to  the  external  angles  of  the  tip 
of  the  horn 2 

2.  Postapical  carina  of  pronotum  sharp  and  slightly  undercut;  male  with  short, 

distinct  carina  back  of  clypeal  horn  arcuate  anteriorly,  in  same  direction 
as  clypeal  horn;  female  smoother  back  of  corresponding  carina  than  in 
front  of  it;  small  shining  species;  8  to  10  mm.     (Baja  California) . 

parvulus  new  species 
Postapical  carina  of  pronotum  rounded;  male  without  carina  or  with  faint 
indication  of  two  low  obscure  tubercles  back  of  clypeal  horn;  female  equally 
rough  behind  and  in  front  of  carina;  usually  dull  or  only  moderately  shining 
(excepting  some  very  large  specimens,  which  may  represent  a  subspecies) ; 
larger  species,  11  to  18  mm schaefferi  (Boucomont) 

Bolborhombus  angulus  (Robinson),  new  combination 

Figure  14,  6 

Bolboceras  angulus  Robinson,  Trans.  Amer.  Ent.  Soc,  vol.  73,  p.  170,  1947. 

Bolborhombus  angulus  (Robinson)  is  a  small  rufotestaceous  species, 
7  to  11  mm.  in  length,  readily  recognized  by  the  form  of  the  head  and 
clypeus.  The  wide  straight  anterior  edge  of  the  clypeus  is  strongly 
carinately  margined.  Behind  this,  from  side  to  side  and  almost  in 
line  with  the  anterior  edges  of  the  ocular  canthi,  is  a  moderately  high 
anteriorly  arcuate  ridge  or  horn,  and  still  farther  back  a  lower  poste- 
riorly arcuate  sharp  carina.  The  median  horn  tends  to  become  some- 
what nodose  at  the  ends  and  is  approximately  midway  between  the 
posterior  carina  and  the  carinate  anterior  edge  of  the  clypeus.  In 
other  characters  angulus  is  very  similar  to  B.  parvulus  and  to  small 
specimens  of  B.  schaefferi  (Boucomont). 

Type. — In  Alark  Robinson  collection,  Philadelphia,  Pa. 


118  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  los 

Type  locality. — Dog  Canyon,  Brewster  County,  Tex. 

Specimens  examined.- — 4,  including  the  holotype. 

Distribution. — Texas:  Dog  Canyon,  Glenn  Spring,  and  Boquillas, 
all  in  Brewster  County.  Arizona:  Miller  Canyon,  Huachuca 
Mountains. 

Season. — July  7  to  29. 

Bolborhombus  parvulus,  new  species 

Figure  14,  a 

HolotjT^pe  male,  length  9  mm.,  width  6  mm.  Shiny,  rufotestaceous, 
densely  hairy  beneath.  Clypeus  extending  obliquel}'^  upward  and 
backward  to  top  of  clypeal  horn;  the  anterior  face  practically  flat, 
its  surface  closely,  very  coarsely  punctate,  with  the  forward  edge 
convex  and  deeply  reticulate-punctate,  the  lateral  edges  parallel  and 
sharply  carmiform;  upper  edge  of  the  horn  shallowly  emarginate,  the 
external  angles  only  slightly  tubcrculiform,  from  them  a  moderately 
high,  sharp  carina  curving  back  each  side  and  ending  somewhat 
abruptly  close  to  the  innermost  edge  of  the  eye;  surface  of  head  back 
of  horn  uneven,  with  close,  niLxed  fine  and  coarse  punctures  and  with  a 
short,  elevated,  transverse,  anteriorly  arcuate  carina  midway  between 
the  similarly  punctate,  sharply  edged,  concave,  depressed  ocular  canthi ; 
the  carina  about  one-third  as  long  as  the  width  of  the  clypeal  horn; 
the  punctures  back  of  the  carina  not  quite  as  close  as  in  front;  the 
occiput  smooth,  closely,  minutely  punctate. 

Pronotum  with  three  rather  deep  foveae  close  behind  the  finely 
margined  anterior  edge,  one  median,  the  others  opposite  mner  eye 
margin  on  each  side;  a  sharp  postapical  carina,  slightly  undercut  on 
posterior  side,  starting  on  each  side  of  the  middle  fovea  and  passing 
close  behind  the  lateral  foveae  toward  but  not  quite  reaching  the 
serrate  side  margin  at  about  anterior  third;  a  second  short,  similar 
carina  on  each  side  straight  back  from  near  the  end  of  the  postapical 
carina  to  the  inner  anterior  edge  of  a  strong  lateral  fovea;  a  shallow, 
coarsely  punctate  groove  half  way  across  the  pronotum  upward  and 
backward  from  each  of  the  tliree  anterior  foveae;  a  row  of  close  coarse 
punctures  in  the  lateral  and  basal  marginal  line,  a  few  m  the  basal 
half  of  median  line,  and  scattered  coarse  punctures  on  both  sides  of 
the  lateral  carma  and  continued  upward  to  opposite  the  humerus,  with 
mixed  fine  and  minute  punctation  throughout;  anterior  angles  slightly 
more  than  a  right  angle;  posterior  angles  obtusely  rounded;  base 
distinctly  margined.  Scutellum  with  mLxed,  very  fine  and  minute 
punctation. 

Elytra  not  margined  basally;  striae  weakly  impressed;  strial  pimc- 
tures  fine;  first  two  striae  interrupted  by  the  scutellum;  intervals 
weakly  convex;  the  surface  minutely  punctate,  very  finely  alutaceous 
under  moderately  high  magnification. 


I 


BOLBOCERATIISri    OF    THE    UNITED    STATES — CARTWRIGHT      119 

Metasternal  plate  twice  as  wide  as  the  lobe  between  the  middle 
coxae,  roughly  punctate,  and  hairy  along  posterior  edges,  Prosternal 
spine  4-cornered,  with  the  anterior  angle  curving  do^vn  to  a  sharp 
point  between  the  coxae  and  with  the  lateral  angles  on  a  transverse 
level  and  the  posterior  curving  do^vnward  to  form  a  sharp  spine. 

Allotype  female,  length  9  mm.,  width  6  mm.  Apparently  identical 
with  the  male  in  every  way  except  in  characters  of  the  head.  Clypeus 
rising  steeply  at  somewhat  less  than  a  right  angle  to  a  sharply  carinate 
edge  seven-eighths  as  wide  and  parallel  to  the  turned-under  anterior 
margin;  the  flat  face  enclosed  being  a  little  less  than  one-third  as  high 
as  long,  then  dropping  sharply  and  gradually  up  again  with  the 
lateral  edges  slightly  diverging  to  the  top  of  the  very  wide  median 
horn,  which  is  widely  and  deeply  emarginate  at  middle,  leaving  the 
two  ends,  or  external  angles,  as  rounded,  thickened,  backward-sloping 
prominences;  back  of  the  emargination,  on  the  same  level,  and  at  a 
point  midway  between  the  ocular  canthi,  is  a  very  short  arcuate 
carina  or  elongate  tubercle,  behind  which  the  general  level  of  the  head 
is  noticeably  lower;  surface  in  front  of  the  horn  deeply,  coarsely 
punctate-reticulate,  between  the  horn  and  the  tubercle  roughly 
coarsely  punctate,  and  behind  the  tubercle  much  smoother,  with 
dispersed,  mixed  fine  and  shallow,  coarse  punctures;  occiput  smooth, 
practically  impunctate. 

Type.' — In  California  Academy  of  Sciences  collection. 

Type  locality.- — Triunfo,  Baja  California,  Mexico.  Holotype  male 
collected  Aug.  7,  1938,  Michelbacher  and  Koss. 

Allotype  and  paratypes. — Allotype  female,  CAS,  and  paratype  male, 
USNM  61656,  San  Venancio,  Baja  California,  Mexico,  Oct.  8,  1941, 
Ross  and  Bohart;  paratype  female,  USNM  61656,  Santa  Rosa,  Baja 
California,  Mexico,  Charles  Schaeffer  collection. 

Remarks.- — Bolborhombus  parvulus  is  very  near  B.  angulus  (Robin- 
son) and  is  about  the  same  size.  The  four  specimens  vary  in  length 
from  8)^  mm.  for  the  smaller  male  to  lOji  mm.  for  the  larger  female. 
The  anterior  edge  of  the  clypeus,  described  as  margined  in  angulus,  is 
almost  exactly  the  same  as  in  the  female  of  parvulus. 

Bolborhombus  schaefferi  (Boucomont),  new  combination 

Figure  14,  c 

Bradycinetus  carinatus  Schaeffer,  Trans.  Amer.  Ent.  Soc,  vol.  32,  p.  251,  1906. 
Bolboceras  schaefferi  Boucomont,  Ann.  Soc.  Ent.  France,  vol.  79,  p.  347,  1910. 

Bolborhombus  schaeferi  (Boucomont)  averages  much  larger  than 
B.  parvulus  and  B.  angulus,  measuring  11  to  18  mm.  in  length,  but  it 
is  very  similar  in  all  characters  except  those  of  the  head,  clypeus,  and 
aedeagus.     Males  show  considerable^variation  in  the  clypeal  horn. 


120  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  i03 

In  smaller  specimens  the  clypeus  slopes  upward  and  backward  from 
the  wide  straight  anterior  edge  to  the  narrower  but  moderately  wide 
and  somewhat  anteriorly  arcuate  apex  of  the  horn,  the  external 
angles  of  which  are  thickened  to  slightly  nodose.  In  large  specimens 
the  external  angles  of  the  horn  become  strongly  elevated  tubercles,  in 
some  individuals  wide  apart,  in  others  relatively  close  together.  The 
head  back  of  the  horn  is  densely,  quite  coarsely  punctate,  occasionally 
with  a  short,  obscure,  anteriorly  arcuate  carina  or  very  low  binodose 
tubercle.  In  the  female  the  anterior  edge  of  the  clypeus  is  nearly 
vertical  to  the  sharply  carinate  margin.  Well  back  and  nearly  in  line 
with  the  anterior  edges  of  the  ocular  canthi  are  two  widely  separated, 
conical  tubercles  with  a  fine  anteriorly  angulate  carina  between  them 
and  a  low  median  swelling  or  tubercle  close  behind  them.  The 
tubercles  are  usually  strongly  developed  but  occasionally  in  small 
specimens  may  appear  only  as  slight  thickenings  of  the  ends  of  the 
angulate  carina. 

Type. — ^Lectotype  male,  present  selection,  USNM  42569. 

Type  locality. — Palmerlee,  Cochise  County,  Ariz. 

Specimens  examined. — 104. 

Distribution. — Arizona:  Cave  Creek  and  Pinery  Canyon  (Chira- 
cahua  Mountains,  Cochise  County),  Dewey,  Douglas,  Globe,  Hua- 
chuca  Mountains,  Fort  Huachuca,  Kits  Peak  (Rincon,  Baboquivari 
Mountains),  Nogales,  *Oracle,  Palmerlee,  Patagonia,  Prescott, 
Reddington,  Madera  Canyon  (Santa  Rita  Mountains),  and  Tucson. 
Texas:  *Alpine,  El  Paso,  *Fort  Davis,  Kingsville,  and  Round  Moun- 
tain. Mexico:  Alamos  (Sonora),  Venedio  (Sinaloa),  Oaxaca  (Oaxaca), 
Santa  Rosa  (Baja  California),  Naco  (Sonora),  Tlahualilo,  and  Rio 
Mayo. 

Season. — July  1  to  October  10,  with  largest  numbers  in  July  and 
August. 


B.  S.  90VIRNMENT  PRINTIH6  OFFICBi  1*13 


U.    S.     NATIONAL    MUSEUM 


PROCEEDINGS.    VOL.    103.    PLATE   3 


•'"^f^rV 


^, 


Frcint  and  dorsal  views  of  three  species  of  Bradycinelulus:  Left,  B.  fossatus  (Haldeman); 
middle,  B.  rex,  new  species;  right,  B.  ferrugineus  (Palisot  de  Bcauvois). 


U.    S.    NATIONAL    MUSEUM 


PROCEEDINGS,    VOL.    103.   PLATE  4 


Differences  in  the  cl\'peai  horn  in  Bolbocerastes:  Upper  left,  B.  serratus  (LeConle);  upper 
right,  B.  regalis,  new  species;  lower  left,  B.  impenalis,  new  species;  lower  right,  B. 
peninsularis  (Schaeffer). 


PROCEEDINGS   OF   THE   UNITED   STATES  NATIONAL   MUSEUM 


byihe 


SMITHSONIAN  INSTITUTION 

U.  S.  NATIONAL  MUSEUM 


Vol.    103  Washington:  1953  No.  3319 

THE  CHRYSOMELID  BEETLES  OF  THE  GENUS 
STRABALA  CHEVROLAT 


By  Doris  Holmes  Blake 


Introduction 


The  first  species  of  this  chrysomehd  genus  was  described  by  IlHger  * 
in  1807  as  Haltica  ruja  from  Pennsylvania,  Melsheimer  collector. 
A  year  later,  Olivier  ^  described  Altica  ferruginea  from  Santo  Do- 
mingo, Palisot  de  Beauvois  collector,  and  Altica  scutellaris  from 
southern  France.  Of  this  latter  locality  von  Harold  ^  said  that  there 
was  no  Altica  colored  anything  like  this  in  southern  France  and  that 
the  locality  was  evidently  an  error.  In  1837,  in  the  Dejean  catalogue, 
Chevrolat  *  listed  under  the  new  generic  name  Strabala  {crpa^aKa, 
having  a  form  thick  and  contracted)  six  specific  names  of  Dejean 
without  descriptions  or  references.  Four  of  them  are  pure  nomina 
nuda;  for  the  two  others,  S.  scutellata  Dejean  and  S.  dominicensis 
Dejean,  the  names  {Altica)  scutellaris?  Olivier  and  (A.)  ferruginea! 
Olivier,  respectively,  are  given  as  doubtful  synonyms.  The  generic 
name  is  consequently  not  published  here.  Chevrolat  changed  the 
locality  for  scutellata  to  America  Borealis.  Of  these  six  names  only 
one  has  since  been  used;  S.  intermedia,  whose  locality  in  this  catalogue 
was  given  as  Cuba,  was  adopted  by  Jacquelin  Du  Val '  in  1857  for  a 

'  Illiger,  Mag.  Insekt.,  vol.  6,  p.  152,  1807. 

«  Olivier,  Entomologie,  .  .  .  vol.  6,  pp.  697,  699,  1808. 

•  von  Harold,  Ooleopterologische  Hefte,  vol.  14,  p.  20, 1875. 

•  Chevrolat,  in  Dejean,  Catalogue  de  la  collection  de  ColSopteres  .  .  .  ed.  3,  p.  413,  1837. 

•  Jacquelin  Du  Val,  in  Ramon  de  la  Sagra,  Historia  .  .  .  de  la  Isla  de  Cuba  (Spanish  ed.),  vol.  7,  p.  IM, 
1857. 

232989—53  121 


122  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  loe 

Cuban  species.  In  d'Orbigny's  Dictionnaire,  Chevrolat'  in  1848 
listed  scutellaris  and  jerruginea  under  Strabala  without  the  question 
marks,  as  well  as  the  four  nomina  nuda  he  had  given  in  1837,  thus 
establishing  the  genus. 

In  1868  Suffrian '  described  Haltica  amhulans  from  Cuba  which  he 
said  belonged  to  Chevrolat's  genus  Strabala,  a  genus  that  Erichson  ' 
had  earlier  merged  with  Lactica.  Suffrian  was  not  convinced,  however, 
that  these  specimens  having  so  poorly  developed  a  basal  sulcus  on  the 
prothorax  really  belonged  to  the  genus  Lactica.  He  stated  that 
although  there  was  an  ill-defined  basal  sulcus,  the  beetles  were  smaller 
than  most  species  of  Altica  and  proportionately  broader,  and  that  the 
color  was  not  blue  but  a  deep  reddish  brown.  This  Cuban  species  that 
he  had  did  not  entirely  correspond  in  coloring  to  Olivier's  description 
of  scutellaris  or  to  Jacquelin  Du  Val's  intermedia,  both  of  which  were 
paler  beneath  than  the  dark  colored  undersm-face  of  his  beetles. 

Boheman  °  described  under  the  genus  Strabala,  two  species,  nigriceps 
from  Buenos  Aires  and  languida  from  Java,  both  of  which  have  been 
referred  to  Lactica.  Although  I  have  not  examined  the  types,  the 
description  of  nigriceps  as  having  the  anterior  angles  of  the  prothorax 
truncate  seems  to  exclude  it  from  Strabala,  and  the  description  of 
languida  is  quite  different  from  that  of  any  of  the  group.  Fall,^"  in 
describing  Altica  testacea  from  Texas,  compared  it  with  rufa,  noting 
that  "rufa  is  much  broader  and  less  convex  and  has  black  legs  and 
antennae."  Fall's  species,  also,  does  not  belong  in  Strabala.  It  differs 
in  general  shape,  the  prothorax  has  quite  different  anterior  angles, 
and  the  scutellum  is  small  and  inconspicuous. 

Jacoby,^^  who  had  put  scutellaris  under  Lactica,  in  the  Supplement 
says  of  Horn,  "Dr.  Horn  remarks  that  Lactica  scutellaris  would  be 
better  placed  in  the  genus  Haltica  and  that  it  is  congeneric  (and  prob- 
ably also  specifically  identical)  with  Haltica  rufa  Uliger."  But 
Jacoby  "could  not  agree  with  this  opinion"  and  kept  scutellaris  under 
Lactica. 

The  British  Museum  (Natural  History)  specimens,  therefore,  are 
under  Lactica,  and  in  the  U.  S.  National  Museum,  following  Horn, 
they  are  under  Altica.  Henshaw  ^^  lists  rufa  under  Disonycha. 
Gemminger  and  Harold  ^^  list  rufa  under  Disonycha  and  scutellaris 
under  Lactica.    Heikertinger,'*  in  the  Junk  catalogue,  nicely  balances 

«  OhevTolat,  in  d'Orbigny,  Dictionnaire  universe!  d'histoire  naturelle,  vol.  12,  p.  52,  1848. 
'  Sufirian,  Arch.  Naturg.,  vol.  34,  p.  182,  1868. 
»  Erichson,  Arch.  Naturg.,  vol.  13,  p.  173,  1847. 

•  Boheman,  Kongliga  Svenska  Fregatten  Eugenies  Resa  .  .  .  ,  vol.  2,  Zoologi,  pt.  1,  Insects,  pp.  189,  190, 
1850. 
M  Fall,  Trans.  Amer.  Ent.  Soc,  vol.  36,  p.  157, 1910. 

"  Jacoby,  Biologia  Oentrali-Americana,  Coleoptera,  vol.  6,  pt.  1,  p.  273, 1884;  Supplement,  p.  259, 1891. 
w  Henshaw,  List  of  the  Coleoptera  of  America,  north  of  Mexico,  p.  112, 1885. 
>'  Gemminger  and  von  Harold,  Oatalogus  coleopterorum  .  .  .,  vol.  12,  p.  3497, 1876. 
M  Heikertinger,  Coleopterorum  catalogus,  pars  166,  pp.  241,  259, 1939. 


CHRYSOMELID    GENUS    STRABALA    CHEVROLAT — BLAKE         123 

the  dispute  by  synonymizing  scutellaris  and  ambulans  with  rufa  and 
placing  them  under  Haltica  and  at  the  same  time  placing  ferruginea 
and  intermedia  as  separate  species  under  Lactica.  Mr.  G.  E.  Bryant 
pointed  out  to  me  the  specimens  labeled  Lactica  scutellaris  in  the 
British  Museum  and  said,  "What  are  these  doing  here?"  I  agreed 
that  it  was  high  time  to  restore  the  original  Chevrolat  generic  name 
Strabala  for  this  group  of  very  similarly  colored  species  that  no  one 
has  Itnown  exactly  what  to  do  with.  On  my  return  to  America,  after 
a  considerable  search  for  the  specimens  of  this  species  that  had  long 
been  removed  from  the  regular  collection,  I  found  that  H.  S.  Barber 
had  also  labeled  them  Strabala. 

The  species  of  this  genus  differ  in  color  among  themselves  little  more 
than  do  those  of  the  genus  Altica,  but  instead  of  being  metallic  blue 
they  are  a  deep  reddish  or  orange  brown  with  dark  or  partly  dark  legs, 
dark  scutellum  (usually) ,  dark  antennae,  and  more  or  less  dark  under- 
surface.  Like  Altica  they  have  no  spots,  vittae,  or  other  markings; 
and  the  variation  in  the  dark  coloration  is  slight.  In  practically  all 
the  North  American  specimens  from  Massachusetts  to  Panamd,  the 
amount  of  dark  coloration  is  very  much  the  same,  that  is,  the  under- 
surface  and  femora  are  only  partly  dark.  Hence  the  tendency  has 
been  to  follow  Horn/^  who  wrote  that  ''Haltica  rufa  occurs  from 
Massachusetts  to  Illinois,  Florida  and  Texas,  extending  through 
Mexico  to  South  America."  There  are,  however,  slight  differences  in 
the  extent  of  the  dark  coloration,  in  the  punctation,  and  in  the  shape 
of  the  aedeagus.  In  Eastern  United  States  from  New  Hampshire  to 
the  Gulf  and  west  to  the  Mississippi,  Louisiana,  and  northern  Texas, 
the  species  known  as  ruja  Illiger  presents  little  variation.  It  is  a  con- 
spicuously punctate,  deep  reddish  brown  species.  In  Florida  occurs  a 
race  that,  in  dried  specimens,  at  least,  is  always  a  pale  yellow-brown, 
in  contrast  to  the  reddish  color  of  rufa.  It  has  also  a  distinctly  aluta- 
ceous  surface  and  is  less  distinctly  punctate  than  rufa.  In  the  speci- 
mens from  Brownsville,  Texas,  to  Panamd,  which  are  shining  and 
almost  impunctate,  in  our  limited  collections  are  two  groups  with 
entirely  differently  shaped  aedeagi — those  that  are  narrowed  at  the 
tip  and  those  with  a  wide,  rounded  tip.  From  the  external  appearance 
I  cannot  distinguish  the  beetles  at  all.  H.  S.  Barber  has  labeled  one 
lot,  with  round-tipped  aedeagi,  as  a  new  species  and  the  other  as 
scutellaris.  It  seems  to  me  that  the  name  scutellaris  cannot  be  applied 
to  any  species  of  Strabala  with  any  certainty.  To  begin  with,  Olivier's 
locality  of  southern  France,  apparently  erroneous,  leaves  us  without 
any  type  locality  in  the  western  hemisphere.  Species  corresponding  in 
coloration  to  Olivier's  description  occur  in  Cuba  and  Central  America 
as  well  as  North  America.     Therefore  I  propose  to  drop  the  name 

W  Horn,  Trans.  Am.  Ent.  Soc,  vol.  16,  p.  232, 1889. 


124  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  i03 

scutellaris  entirely,  as  retaining  it  would  only  promote  confusion. 
Among  the  specimens  from  Central  America  with  the  aedeagus  nar- 
rowed at  the  tip  there  is  a  wide  variation  in  the  acuteness  and  in  the 
width  of  the  tip.  I  think  there  are  probably  a  number  of  species,  but 
since  the  coloring  is  so  nearly  uniform  and  the  variation  in  the 
aedeagi  only  a  matter  of  degree,  I  hesitate  to  give  them  specific  names 
and  am  calling  them  only  subspecies. 

In  the  Bowditch  collection  at  the  Museum  of  Comparative  Zoology 
(MCZ)  are  seven  specimens  from  South  America  (two  from  Venezuela 
and  the  rest  from  Colombia)  that  are  uniformly  somewhat  larger  than 
the  others  of  the  genus  and,  unlike  the  North  and  Central  American 
specimens,  are  dark  beneath,  as  in  the  majority  of  the  West  Indian 
specimens.  In  the  British  Museum  is  a  series  from  Trinidad,  col- 
lected by  G.  E.  Bryant,  also  dark  beneath  but  smaller,  paler,  and  with 
a  more  rounded  tip  to  the  aedeagus. 

In  the  West  Indies  the  prevailing  color  of  the  legs  and  undersurface 
of  the  specimens  is  entirely  dark,  but  in  one  species,  at  least,  the  pos- 
terior femora  are  partly  reddish,  as  in  the  continental  species.  In 
another  species  from  Hispaniola,  the  scutellum  is  not  black  but 
reddish  lil^e  the  elytra,  the  only  one  so  colored  that  I  have  examined. 
Jacoby  wrote  of  specimens  with  a  red  scuteUum  from  Juquila,  Mexico, 
of  which  he  had  three.  One  specimen  from  Juquila  in  his  collection 
is  a  yellow  Lactica  with  a  pronounced  basal  sulcus  on  the  prothorax, 
typical  of  Lactica,  and  a  pale  scutellum.  This  may  not  be  the  one  to 
which  he  referred.  These  slight  differences  in  coloration  of  the  speci- 
mens from  the  different  islands  of  the  West  Indies  are  accompanied 
by  a  slightly  different  aedeagus.  Still,  as  in  the  case  of  the  Central 
American  specimens,  I  hesitate  to  give  them  specific  names  and  am 
calling  them  only  subspecies.  Much  more  work  should  be  done  on 
the  group  and  many  more  specimens  should  be  studied.  Here  I 
attempt  to  assemble  the  species  that  in  the  catalogues  are  divided 
among  two  or  three  genera  and  to  call  attention  to  their  points  of 
difference. 

The  group  is  closer  to  Disonycha  than  to  either  Altica  or  Lactica. 
The  head,  while  similar  to  Disonycha,  has  the  circle  of  punctures 
nearer  to  the  tubercles  and  the  carina  shorter  and  broader  than  in 
most  species  of  Disonycha.  The  antennal  joints  are  similar  in  being 
robust  and  rather  short,  but  differ  in  having  the  third  joint  a  little 
longer  than  the  fourth,  instead  of  the  fourth  longer  than  the  third. 
The  prothorax  is  much  like  that  of  Disonycha  except  that  below  the 
apical  angle  there  is  a  tiny  V-shaped  notch  which  does  not  occur  in 
Disonycha.  The  basal  depression  over  the  scuteUum,  often  found  in 
Disonycha  too,  is  longer  and  more  pronounced  in  Strabala.  The 
elytra  are  distinctive  in  being  more  convex  and  oval  than  in  Lactica, 
Altica,  or  Disonycha. 


CHRYSOMELID  GENUS  STRABALA  CHEVROLAT — BLAKE         125 


^.S.Qmbulans(5uffrloi)  A.S.Qmbubos  »sp JamalcsnatS  ■.  S  ferru(JIoe<lK)iwier)  y.S  ambulons  ssp. poerforicens(s 


k.  Scolumblana 


/.  S  lnfcrmed{Q  Jorq.duVat 


■.   StrinltalO 


Figure  17. — Species  of  Strabala  Chevrolat:  a,  rufa  (Illiger);  i,  rufafloridana,  new  subspecies; 
c, acuminata,  new  species;  d,  rotunda,  new  species;  e,  acuminata  teapensis,  new  sub- 
species /,  acuminata  costaricensis,  new  subspecies;  g,  ambulans  (Suffrian);  h,  ambulant 
jamaicensis,  new  subspecies;  i,  ferruginea  (Olivier);  /,  ambulans  puertoricensis,  new 
subspecies;  k,  columbiana,  new  species;  /,  intermedia  Jacquelin  Du  Val;  m,  trinitatis, 
new  species. 


126  PROCEEDESrGS    OF    THE    NATIONAL   MUSEUM  vol.  io3 

Genus  Strabala  Chevrolat 

Description. — Between  3.5  and  6  mm.  in  length,  oval,  deep  yellowish 
or  orange  brown  or  reddish  brown,  with  antennae  dark,  legs  and  under- 
surface  more  or  less  dark,  and  scutellum  usually  conspicuously  dark; 
moderately  shining,  most  species  very  minutely  and  confusedly 
punctate,  only  the  North  American  species,  iS.  rufa  (Illiger),  very 
distinctly  punctate. 

Head,  except  for  mouth  parts  that  are  usually  a  little  darker,  deep 
yellowish  or  reddish,  eyes  entire  and  widely  separated,  the  interocular 
space  being  more  than  half  the  width  of  the  head;  occiput  smoothly 
rounded  down  to  tubercles,  with  scattered  and  not  very  dense  or  coarse 
punctures;  at  the  base  of  occiput,  not  visible  except  when  the  head  is 
a  little  protruded,  a  groove  running  from  behind  one  eye  to  the  other; 
on  either  side  above  the  frontal  tubercles  and  near  the  eye  a  depression 
consisting  of  a  large  fovea,  or  more  frequently  a  circle  of  punctures; 
frontal  tubercles  plainly  marked,  a  broadly  rounded  and  somewhat 
produced  area  between  antennal  sockets  that  extends  and  widens  down 
the  front  nearly  to  the  labrum,  area  under  sockets  a  little  depressed 
and  on  sides  below  eyes  the  cheeks  rounded  out  and  densely  punctate. 
Antennae  rather  stout,  not  quite  reaching  the  middle  of  the  elytra, 
first  two  joints  swollen,  usually  paler,  remainder  piceous,  thh-d  joint 
a  little  longer  than  fourth,  fourth  and  fifth  about  equal.  Prothorax 
not  quite  twice  as  wide  as  long,  with  rounded  sides  and  narrow  margin, 
the  apical  angle  resembling  that  of  Disonycha  except  that  it  is  sepa- 
rated from  the  rest  of  the  margin  by  a  distinct  though  tiny  V-shaped 
indentation,  basal  angle  with  a  small  tooth  and  obliquely  cut,  disk 
smooth  and  rather  flat,  with  more  or  less  of  a  depression  along  base 
and  not  distinctly  limited  at  the  ends  as  m  Lactica.  Scutellum  large, 
usually  conspicuously  dark,  contrasting  with  the  reddish  elytra. 
Elytra  broader  than  prothorax,  moderately  convex,  with  small 
humeri  and  narrow  margin,  not  depressed ;  surface  usually  smooth,  not 
distinctly  punctate  except  in  S.  rufa,  entirely  yellowish  or  reddish 
brown.  Epipleura  broad,  gradually  diminishing  and  not  quite  reach- 
ing the  apex.  Body  beneath  variable  in  color,  the  prosternum  red- 
dish, the  breast  and  abdomen  and  legs  in  West  Indian  species  usually 
but  not  always  dark,  in  North  American  species  the  breast  dark  on 
the  sides,  the  femora  pale  except  in  apical  half  or  sometimes  entirely 
pale,  the  tibiae  and  tarsi  dark,  the  sides  and  apex  of  the  abdomen 
sometimes  also  dark;  in  Trinidad  and  South  American  species,  the 
coloring  beneath  dark  as  in  the  West  Indian  specimens.  Anterior 
coxal  cavities  open,  front  coxae  narrowly  separated,  legs  rather  short, 
hind  femora  moderately  thickened,  the  tibiae  as  in  Disonycha  with  a 
slight  ridge,  a  spur  at  the  end  of  posterior  tibiae,  claws  appendiculate. 


CHRYSOMELID    GENUS    STRABALA    CHEVROLAT — BLAKE         127 

Aedeagus  a  simple,  curved  structure  with  the  tip  varying  in  shape 
from  acute  to  truncate  and  broadly  rounded. 

Genotype. — Strahala  ferruginea  (Olivier),  present  designation. 

Key  to  the  species  of  Strabala 

1.  Scutellum  reddish  (Hispaniola) ferruginea   (Olivier) 

Scutellum  dark  brown  or  piceous 2 

2.  Species  from  the  West  Indies  exclusive  of  Trinidad 3 

Species  from  North  America 5 

Species  from  South  America  and  Trinidad 9 

3.  Body  beneath  more  or  less  ferrugineous,  hind  femora  ferrugineous  at  base, 

dark  at  apex  (Cuba) intermedia  Jacquelin  Du  Val 

Body  beneath  except  prosternum  more  or  less  dark,  hind  femora  entirely 
dark 4 

4.  Aedeagus  tapering  gradually  to  a  rounded  tip  (Cuba) ambulans  (Suffrian) 

Aedeagus  tapering  gradually  to  a  more  acute  but  still  rounded  tip  (Jamaica). 

ambulans  jamaicensis,  new  subspecies 
Aedeagus  distinctly  truncate  at  tip  (Puerto  Rico). 

ambulans  puertoricensis,  new  subspecies 

5.  Elytra  punctate  or  alutaceous  or  both  (North  of  Mexico) 6 

Elytra  not  distinctly  punctate  or  alutaceous  (Texas  and  southward) 7 

6.  Elytra  deep  reddish  brown,  distinctly  punctate  (New  Hampshire  south  and 

westward  to  the  Mississippi,  Louisiana,  Texas,  and  along  the  Gulf  to  Florida). 

rufa  (lUiger) 

Elytra    (in    dried    specimens)    yellow,    less   distinctly    punctate,    alutaceous 

(Florida) rufa  floridana,  new  subspecies 

7.  Aedeagus  well  rounded  at  tip  and  with  a  slight  constriction  behind   the   tip 

(Mexico  to  Panamd) rotunda,  new  species 

Aedeagus  gradually  narrowing  towards  the  tip,  not  constricted  behind  the 
tip 8 

8.  Aedeagus  narrowed  to  a  rounded  tip,  hind  femora  dark  at  apex  (Texas) . 

acuminata,  new  species 
Aedeagus  narrowed  to  a  truncate  tip,  hind  femora  entirely   reddish   brown 

( Mexico) acuminata  teapensis,  new  subspecies 

Aedeagus  narrowed  to  a  broader  truncate  tip,  hind  femora  varying  from 
entirely  reddish  to  reddish  with  a  dark  apex  (Guatemala,  Costa  Rica) . 

acuminata  costaricensis,  new  subspecies 

9.  Large,  6  to  6  mm.,  aedeagus  with  an  acute  tip  (Colombia,  Venezuela). 

colombiana,  new  species 
Smaller,  4  to  5  mm.,  aedeagus  with  a  somewhat  truncate  tip  (Trinidad). 

trinitatis,  new  species 


Strabala  rufa  (Illiger) 

Figure  17,  a 

Haliica  rufa  Illiger,  Mag.  Insekt.,  vol.  6,  p.  152,  1807, 

Illiger's   description  of  the  thickly  punctate  elytra  distinguishes 
this  species  from  the  others  of  the  genus.    The  coloration  is  usually 


128  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  103 

deep  reddish  brown  above,  with  entirely  dark  antennae,  dark  scutel- 
lum,  often  entirely  dark  anterior  legs,  dark  apical  end  of  the  posterior 
femora,  dark  breast,  and  sometimes  also  dark  abdomen.  Throughout 
its  range  over  the  country  east  of  the  Mississippi  and  in  Louisiana  and 
Texas,  there  seems  to  be  very  little  variation  except  for  the  paler, 
less  distinctly  punctate  race  that  occurs  in  Florida.  The  type  locality 
is  Pennsylvania,  collected  by  Melsheimer. 

Distribution. — New  Hampshire:  Hampton,  S.  A.  Shaw.  Massachu- 
setts: Holyoke,  Chicopee,  F.  Knab.  Connecticut:  Hartford,  F.  Knab. 
Rhode  Island:  Watch  Hill,  W.  Robinson.  New  York:  West  Point, 
W.  Robinson.  New  Jersey:  Boynton.  Illinois:  C.  Thomas.  Mich- 
igan: Detroit,  Hubbard  and  Schwarz.  Maryland.  Washington, 
D.  C.  Virginia:  Fredericksburg,  Clifton,  J.  C.  Bridwell;  Nelson 
County,  W.  Robinson;  Stone  Creek,  Lee  County,  Hubbard  and 
Schwarz;  Vienna,  J.  C.  Bridwell;  West  Falls  Church,  E.  A.  Chapin. 
West  Virginia:  Aurora.  Kentucky:  H.Soltau.  Tennessee:  Memphis, 
H.  Soltau.  North  Carolina:  Graybeard  Mountain.  South  Carolina: 
Florence.  Georgia:  Savannah.  Alabama:  Coleta,  H.  M.  Smith. 
Mississippi:  Greenwood,  T.  H.  Parks.  Louisiana:  Baton  Rouge, 
T.  H.  Jones;  Port  Allen,  J.  L.  Lauderdale;  New  Orleans,  H.  Soltau. 
Texas:  Liberty,  R.  A.  Vickery.  Two  specimens,  both  females, 
USNM  (Brooklyn  Museum  collection),  labeled  simply  "Mexico,"  are 
apparently  this  species. 

Strabala  riifa  floridana,  new  subspecies 

Figure  17,  6 

This  race  differs  from  the  reddish  bro\\Ti  one  found  elsewhere  in 
eastern  United  States  by  being  (in  dried  specimens)  pale  yellow-brown 
with  the  dark  markings  as  in  typical  rufa.  The  elytra  are  less  dis- 
tinctly punctate  than  in  rufa  and  distinctly  alutaceous. 

Tyjpe. — Holotype  male  and  7  paratypes,  USNM  61202,  labeled 
simply  "Fla.,"  collector  F.  Knab. 

Distribution. — Florida:  Lake  Ashby,  Capron,  Indian  River,  all 
collected  by  Hubbard  and  Schwarz;  Lake  Wales,  Lakeland,  E.  M, 
Craighead;  Jacksonville,  Ashmead;  Sanford,  Van  Duzee;  St.  Peters- 
burg (on  sweet  corn) ;  Ft.  Myers  (on  sweet  and  white  potato) ;  Oneco 
(on  spinach);  Homestead  (on  string  beans);  West  Palm  Beach  (on 
sweet  potato  and  lima  beans) . 

Strabala  acuminata,  uew  species 

Figure  17,  c 

From  4.5  to  5.5  mm.  in  length,  oval,  shining,  orange-brown,  with 
dark  antennae,  dark  scutellum,  darkened  sides  to  breast,  and  dark 
anterior  legs  and  apical  part  of  posterior  femora,  tibiae,  and  tarsi. 


CHRYSOMELID    GEsSTUS    STRABALA    CHEVROLAT — BLAKE         129 

Head  shiny,  only  a  few  scattered  punctures  over  occiput  besides 
the  fovea  or  circle  of  depressed  punctures  near  the  eye;  mouth  parts 
frequently  edged  with  deeper  brown.  Antennae  deep  brown,  with 
the  three  basal  joints  frequently  paler.  Prothorax  shiny,  almost 
impunctate,  only  under  high  magnification  are  very  fine  punctures 
visible ;  basal  sulcus  usually  not  very  long  or  pronounced,  but  somewhat 
variable  in  different  specimens,  as  is  usual  in  the  genus.  Scutellum 
dark.  Elytra  mirror  smooth,  shining,  impunctate,  sometimes  very 
indistinctly  alutaceous.  Body  beneath  with  sides  of  breast,  anterior 
legs,  apical  part  of  posterior  femora,  tibiae,  and  tarsi  dark.  Length 
4.4  to  5.5  mm.;  width  2.  4  to  3  mm. 

Type.— Holotype  male  and  2  paratypes,  USNM  61203  (Brooklyn 
Museum  collection),  collected  at  the  Esperanza  Ranch,  Brownsville, 
Texas,  August  18  (no  year  given). 

Other  localities. — Texas:  Brownsville,  R.  A.  Vickery;  another  speci- 
men, from  same  locality  (on  string  bean  leaves);  San  Benito  (on 
bean  foliage);  Hidalgo  (on  black-eyed-pea  leaf). 

Remarks. — The  shiny,  almost  impunctate  surface  with  little  evidence 
of  alutaceous  dullness  at  once  differentiates  this  from  Strabala  rufa 
(lUiger).  There  are  no  records  of  it  farther  north  than  the  Brownsville 
region. 

Strabala  acuminata  teapensis,  new  subspecies 

Figure  17,  e 

Lactica  scutellaris  Olivier,  Jacoby   (part),   Biologia  Centrali- Americana,  vol.  6, 
pt.  1,  p.  273,  1884. 

This  differs  from  the  Brownsville,  Texas,  race  in  having  less  dark 
coloring,  the  femora  all  being  deep  reddish,  the  tibiae  and  tarsi  only 
dark,  and  the  sides  of  the  breast  not  so  dark.  The  upper  surface  is 
a  shining  deep  reddish  brown.  The  aedeagus  has  a  somewhat  trim- 
cate  tip. 

Type. — Holotype  male,  in  the  British  Museum  (Natiu-al  History), 
from  the  Biologia  Centrali-Americana  material,  labeled  by  Jacoby 
Lactica  scutellatis  Olivier,  from  Teapa,  Tabasco,  Mexico,  collected  in 
March  by  H.  H.  Smith;  1  paratype  in  U.  S.  National  Museum, 
USNM  61204;  and  2  paratypes  in  Museum  of  Comparative  Zoology 
from  same  series. 

Strabala  acuminata  costaricensis,  new  subspecies 

Figure  17,  / 

Lactica  scutellaris  Olivier,  Jacoby  (part),  Biologia  Centrali-Americana,  vol.  6, 
pt.  1,  p.  273,  1884. 

This  differs  from  the  Brownsville,  Texas,  and  Teapa,  Mexico,  races 
by  having  a  still  wider  truncate  tip  to  the  aedeagus.     In  coloring,  it 


130  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  i03 

is  somewhat  deeper  red  than  the  Brownsville  specimens  and  differs 
from  the  Teapa  specimens  in  having  the  anterior  femora  and  apical 
part  of  the  posterior  femora  as  well  as  the  sides  of  the  breast  dark. 

Type. — Holotype  male  and  1  paratype,  USNAl  61205,  collected  at 
Vara  Blanca,  Costa  Rica,  1,700  meters,  July  7,  1928,  by  F.  Never- 
mann;  1  paratype  in  British  Museum  (Natural  History). 

Other  localities. — Costa  Rica:  San  Jose,  1,000-2,000  meters,  Aug.  25, 
1928,  F.  Nevermann.  Guatemala:  Yepocapa,  August  1948,  H.  T. 
Dalmat;  Cob^n,  Alta  Vera  Paz,  May  1926,  J.  M.  Aldrich;  Purula, 
Champion 

Strabala  rotunda,  new  species 

Figure  17,  d 

Lactica  scutellaris  Olivier,  Jacoby   (part),  Biologia  Centrali- Americana,  vol.  6, 
pt.  1,  p.  273,  1884. 

From  4  to  5.5  mm.  in  length,  oval,  shining  although  faintly  alu- 
taceous,  very  finely  punctate,  yellowish  or  reddish  brown  with  dark 
antennae,  dark  apex  to  femora,  dark  tibiae  and  tarsi,  and  the  breast 
and  abdomen  often  darkened  on  the  sides,  varying  to  entirely  dark 
(rare) . 

Head  reddish,  with  the  mouth  parts  usually  deeper  brown;  finely 
punctate  over  occiput,  with  the  usual  fovea  or  circle  of  depressed 
punctures.  Pro  thorax  shiny  and  very  finely  punctate  with  a  sulcus 
along  the  base  well  developed.  Scutellum  dark.  Elytra  faintly 
alutaceous  and  finely  punctate.  Body  beneath  varying  in  color 
from  having  the  sides  of  breast  and  abdomen  deeper  brown  to  being 
entirely  dark,  the  latter  rare ;  femora  with  apical  half  dark,  tibiae  and 
tarsi  dark.     Length  4  to  5.4  mm.;  width  2.6  to  3  mm. 

Type. — Holotype  male,  USNM  61206,  and  3  paratypes  in  British 
Museum  (Natural  History)  collected  at  Volc^n  de  ChiriquI,  Panamd, 
by  Champion. 

Other  localities. — Panamd:  Panama,  Aug.  13,  1946,  by  N.  L.  H. 
Kxauss;  Almirante,  Sept.  4,  1938,  H.  Dybas;  Bella  Vista,  Aug.  7,  1924, 
N.  Banks;  Mindi  Dairy  (on  para  grass),  Mindi,  Canal  Zone,  July 
1918,  H.  F.  Dietz  and  J.  Zetek;  Plantation  Borracho,  Canal  Zone 
(swept  from  grass  under  papaya  trees),  July  1918,  H.  F.  Dietz  and 
J.  Zetek;  Bugaba,  Champion  (in  British  Museum).  Guatemala: 
Trece  Aguas,  Alta  Vera  Paz,  May  (date  not  given),  H.  S.  Barber  and 
E.  A.  Schwarz.  Mexico:  In  bananas  from  Mexico  taken  at  Mobile, 
Alabama,  July  19,  1929;  with  orchids  from  Guerrero  (State),  taken  at 
Laredo,  Texas,  Feb.  28,  1940;  with  orchids  from  Tamazunchale, 
San  Luis  Potosi,  taken  at  Laredo,  Texas,  Feb.  28, 1940;  with  tomatoes 
from  El  Mante,  Tamaulipas,  taken  at  El  Paso,  Aug.  1,  1947;  Colima, 


CHRYSOMELID    GENUS    STRABALA    CHEVROLAT BLAKE  131 

Hoge  and  Conradt;  Jalapa,  Hoge;  on  green  peppers  from  Manuel, 
Tamaulipas,  taken  at  Laredo,  Texas,  Jan.  7,  1935. 

Remarks. — H.  S.  Barber  has  attached  a  specific  name  from  the 
Mexican  locaUties  to  this  species,  but  since  the  range  is  as  far  south 
as  Panamd,  his  name  would  be  somewhat  misleading.  The  rounding 
of  the  tip  of  the  aedeagus  varies  considerably  but  the  slight  con- 
striction behind  the  tip  is  always  present,  and  sets  the  species  apart 
from  the  rest.  The  only  other  species  with  a  similar  aedeagus  tip 
in  the  group  is  Strabala  intermedia  Jacquelin  Du  Val,  from  Cuba. 
In  all,  15  specimens  of  S.  rotunda  have  been  taken  on  orchids 
from  Mexico  at  the  port  of  entry  to  this  country. 

Strabala  ambulans  (SufiFrian) 

Figure  17,  g 
Haltica  ambulans  Sufifrian,  Arch.  Naturg.,  vol.  34,  p.  182,  1868. 

Specimens  from  Cuba  agree  with  SufFrian's  detailed  description  of 
his  Cuban  H.  ambulans.  The  upper  surface  is  faintly  alutaceous  and 
finely  punctate,  in  color  a  deep  reddish  brown  with  dark  scutellum 
and  dark  legs,  breast,  and  abdomen. 

Distribution. — Cuba:  Cayamas,  E.  A.  Schwarz;  Central  Jaronu, 
L.  C.  Scaramuzza;  Baragua,  L.  C.  Scaramuzza,  L.  D.  Chiistenson 
(MCZ);  upper  Yara  Valley,  Oct.  18,  1928,  L.  C.  Scaramuzza  (MCZ) ; 
Soledad,  Cienfuegos,  Nov.  7,  1926,  P.  J.  Darlington  (MCZ);  Aguada 
(MCZ);  Mountains  north  of  Imias,  eastern  Oriente  Province,  July 
25-28,  3-4,000  feet,  P.  J.  Darhngton  (MCZ). 

Strabala  ambulans  jamaicensis,  new  subspecies 

Figure  17,  h 

Between  3.5  and  4,5  mm.  in  length,  oval,  faintly  shining,  indis- 
tinctly alutaceous,  very  finely  punctate,  reddish  brown,  the  mouth 
parts,  antennae,  legs,  breast,  and  abdomen  dark. 

Head  with  scattered  punctures,  the  usual  fovea  or  circle  of  punc- 
tures between  eye  and  frontal  tubercle  on  each  side.  Antennae  with 
the  basal  joints  a  little  paler.  Prothorax  having  a  distinct  sulcus 
along  the  base.  Scutellum  dark.  Elytra  very  faintly  punctate. 
Body  beneath  with  breast,  abdomen,  and  legs  dark.  Length  3.5  to 
4.5  mm.;  width  2  to  2.3  mm. 

Type. — Holotype  male  and  10  paratypes,  USNM  61207,  collected 
at  Manchioneal,  Jamaica,  Jan.  30,  1939,  by  E.  A.  Chapin  and  R.  E, 
Blackwelder. 

Other  localities. — Jamaica:  Roaring  River  Falls,  St.  Ann's  Parish, 
Sept.  13,  1917,  H.  Morrison;  Fern  Gully,  St.  Ann's  Parish,  Sept.  14, 


132  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  los 

1917,  H.  Morrison;  Mandeville,  Van  Duzee,  A.  E.  Wight  (MCZ); 
Montego  Bay,  Mar.  2,  1911;  Balaclava  (MCZ). 

Remarks. — This  subspecies  differs  only  slightly  from  the  Cuban  race 
in  having  a  more  acute  though  still  rounded  tip  to  the  aedeagus. 

Strabala  ambulans  puertoricensis,  new  subspecies 

Figure  17,  j 

Between  4.5  and  5  mm.  in  length,  oval,  shiny,  elytra  not  alutaceous, 
very  indistinctly  punctate,  reddish  brown,  antennae,  mouth  parts, 
legs,  breast  and  abdomen  dark. 

Head  with  scattered  punctures  and  a  fovea  or  circle  of  punctures 
near  the  eyes.  Antennae  with  paler  basal  joints.  Prothorax  not  at 
all  clearly  depressed  along  the  base.  Scutellum  dark.  Elytra 
quite  shiny,  not  alutaceous,  very  finely  and  indistinctly  punctate. 
Body  beneath  dark  except  the  prosternum.  Length  4.5  to  5  mm.; 
width  2.5  to  3  mm. 

Type. — Holotype  male  and  1  paratype,  USNM  61208;  1  paratype 
in  Museum  of  Comparative  Zoology,  taken  at  Adjuntas,  Puerto  Rico, 
Nov.  1,  1932,  by  R.  G.  Oakley. 

Other  localities. — Puerto  Rico:  Lares,  September  1921,  G.  N.  Wol- 
cott;  Arecibo,  June  17,  1932  (on  eggplant  leaf),  Anderson  and  Mills; 
Mayaguez,  January  1899,  A.  Busck;  Villalba,  June  18,  1934,  C.  M. 
Matos  (MCZ);  Mayaguez,  December  1932,  V.  Alexandrina  (MCZ). 

Remarks. — The  shiny,  not  alutaceous  elytra,  the  larger  size,  and  the 
broad,  almost  truncate  tip  of  the  aedeagus  are  unlike  the  Cuban  and 
Jamaican  races  of  the  species. 

Strabala  ferruginea  (Olivier) 

Figure  17,  i 

Altica  ferruginea  Olivier,  Entomologie  .  .  .  ,  vol.  6,  p.  697,  1808. 
Strabala  ferruginea  Chevrolat,   in    d'Orbigny,    Dictionnaire   universel  d'histoire 
naturelle,  vol.  12,  p.  52,  1848. 

Olivier's  description  of  the  scutellum  of  this  species  as  ferrugineous 
would  differentiate  it  from  the  rest  of  the  West  Indian  species  even  if 
he  had  not  given  the  type  locality  as  Santo  Domingo.  He  described 
the  color  of  the  undersurface  as  black  and  this  agrees  with  many  of 
the  specimens,  but  there  is  a  certain  variation  in  the  coloring,  some 
specimens  being  entirely  ferrugineous  beneath,  others  with  dark  legs 
and  reddish  breast  and  abdomen,  others  with  dark  legs,  breast,  and 
abdomen.  Moreover  the  pronotum  and  elytra  are  finely  and  a  little 
more  distinctly  punctate  than  in  Cuban  and  Jamaican  species  and 
not  at  all  alutaceous.  The  aedeagus,  having  a  truncate  tip,  resembles 
the  Puerto  Rican  specimens  more  than  the  Cuban  or  Jamaican. 


CHRYSOMELID    GENUS    STRABALA    CHEVROLAT — BLAKE         133 

Distribution. — Haiti:  La  Vanneau,  June  1920,  Bizotan,  Nov.  3, 
1925,  Diguini,  Apr.  14,  1925,  all  collected  by  W.  A.  Hoffman;  Grande 
Riviere,  W.  M.  Mann;  He  de  la  Tortue,  April  1929,  E.  C.  and  G.  M. 
Leonard;  Port-au-Prince  and  vicinity,  October  1934,  P.  J.  Darlington 
(MCZ);  Miragoane,  October,  November,  P.  J.  Darlington  (MCZ); 
Carrefour,  May  1-3, 1908,  M.  Cameron  (British  Museum).  Dominican 
Republic:  August  Busck;  Macoris,  Mar.  26, 1913,  P.  G,  Russell;  Puerto 
Plata,  Hurst  (MCZ);  Santiago,  1938,  P.  J.  Darlington  (MCZ) 
Slinchez,  July  1938,  P.  J.  Darlington  (MCZ). 

Strabala  intermedia  Jacquelin  Du  Val 

Figure  17,  I 

Strabala  intermedia  Jacquelin  Du  Val,  in  Ram6n  de  la  Sagra,  Historia  .  .  .  de  la 
Isla  de  Cuba  (Spanish  ed.),  vol.  7,  p.  129,  1857. 

In  coloring,  three  old  specimens  in  the  Museum  of  Comparative 
Zoology  labeled  "Cuba"  resemble  Jacquelin  Du  Val's  short  description. 
They  are  deep  yellowish  brown  with  dark  antennae,  dark  scutellum, 
dark  tibiae  and  tarsi;  the  anterior  femora  and  the  apex  of  the  pos- 
terior femora  are  dark.  In  size  they  are  slightly  larger  than  the 
measurements  given  for  S.  intermedia;  instead  of  being  3%  mm.  they 
are  4.5  to  5.5  mm.  in  length.  In  two,  both  females,  the  basal  thoracic 
sulcus  is  pronounced,  almost  like  that  of  a  true  Lactica,  but  in  the  third 
specimen,  a  male,  the  depression  along  the  base  is  less  marked.  The 
surface  is  faintly  alutaceous  and  iBnely  punctate.  In  aU  three  speci- 
mens the  head  is  rather  densely  punctate  in  addition  to  the  circle 
of  depressed  punctures  near  the  eye.  The  coloring  of  these  specimens 
is  more  like  that  of  the  North  American  species.  The  aedeagus, 
although  essentially  the  same  as  in  all  the  others,  is  distinctive  in 
being  slightly  constricted  behind  the  tip  as  in  the  Mexican  and  Central 
American  species  S.  rotunda,  but  differs  in  having  a  more  acute  tip. 

Strabala  colomhiana,  new  species 

Figure  17,  k 

From  5  to  6  mm.  in  length,  oblong  oval,  deep  reddish  brown,  the 
antennae,  breast,  abdomen,  and  legs  dark,  shining,  not  alutaceous, 
very  finely,  and  the  elytra  rather  densely,  punctate. 

Head  usually  entirely  deep  reddish  brown,  shining,  with  scattered 
fine  punctures  over  occiput  and  the  usual  depressed  circle  of  punctiu-es 
or  fovea  near  eye.  Antennae  short,  dark,  the  two  basal  joints  paler. 
Prothorax  shiny,  finely  punctate,  a  depressed  line  near  base.  Scutel- 
lum dark.  Elytra  shiny,  very  finely  and  indistinctly  punctate.  Body 
beneath  with  breast  and  abdomen  deep  piceous,  legs  dark.  Length 
5  to  6  mm.;  width  2.8  to  3.2  mm. 


134  PROCEEDINGS    OF    THE    NATIONAL    MUSEUM 


Type. — Holotype  male,  in  Museum  of  Comparative  Zoology,  and 
1  para  type,  USNM  61209,  from  Bogota,  Colombia,  from  the  Bow- 
ditch  collection,  Jacoby's  second  collection. 

Other  localities. — Colombia;  Nueva  Granada;  Venezuela;  and  Cara- 
cas; all  in  the  Bowditch  collection,  Jacoby's  second  collection. 

Remarks. — Among  the  material  presented  to  the  U.  S.  National 
Museum  from  the  Bowditch  collection  is  a  single  old  specimen 
of  Strabala  sp.  with  the  label  "Lactica—Strabala  haematina  Dej" 
from  "Nov.  Granad."  This  may  be  one  of  the  specimens  listed  in 
the  Dejean  Catalogue  from  Cartagena  since  it  bears  the  old  Dejean 
Catalogue  Dame.  The  aedeagus  of  this  old  specimen  has  a  rounded 
tip  quite  unlike  the  one  from  Bogota  with  its  acute  tip,  and  is  probably 
not  the  same  species. 

Strabala  trinitatis,  new  species 

Figure  17,  m 

From  4  to  4.9  mm.  in  length,  oval,  shiny,  not  alutaceous,  finely 
but  distinctly  punctate,  yellowish  or  reddish  brown  with  darker 
antennae,  scutellum,  legs,  breast  and  abdomen. 

Head  entirely  reddish,  shiny,  finely  punctate  over  occiput.  Anten- 
nae dark,  with  the  first  two  or  three  joints  paler.  Prothorax  shiny, 
finely  punctate,  with  a  moderately  developed  sulcus  over  the  scutel- 
lum. Elytra  distinctly  and  rather  densely  punctate,  shiny,  not  at 
all  alutaceous.  Body  beneath  with  breast,  abdomen,  and  legs  deep 
reddish  brown.     Length  4  to  4.9  mm.;  width  2.5  to  2.7  mm. 

Type. — Holotype  and  4  paratypes,  British  Museum,  and  1  paratype, 
USNM  61210,  collected  m  Trinidad  m  1903  by  G.  E.  Bryant. 

Other  localities. — Two  specimens  (USNM)  collected  by  A.  Busck  on 
June  12  at  Montserrat,  Trinidad. 

Remarks. — The  specimens  in  this  series,  although  of  the  coloration  of 
the  ones  from  Colombia  and  Venezuela,  are  smaller,  more  oval,  and 
distinctly  though  finely  punctate.  The  tip  of  the  aedeagus  is  truncate 
and  not  at  all  acute  as  in  the  single  male  specimen  from  Bogota. 


B.  S.  GOVERNHENT  PRINTING  OFPICEi  I9SS 


PROCEEDINGS   OF   THE   UNITED   STATES  NATIONAL  MUSEUM 


Issued 


SMITHSONIAN  INSTITUTION 

U.  S.  NATIONAL  MUSEUM 
Vol.   103  Washington:  1953  No.  3320 

AMERICAN  BITING  MIDGES  OF  THE  HELEID  GENUS 

MONOHELEA 


By  Willis  W.  Wirth* 


The  genus  Monohelea  was  founded  by  Eaeffer  (1917)  for  a  new 
species,  hieroglyphica,  from  Paraguay,  and  three  other  American 
species  were  included:  Ceratopogon  seguax  Williston,  1896;  C.  macuLi- 
pennis  CoquLllett,  1905;  and  C.  nebulosa  Coquillett,  1901.  Since  then 
a  number  of  species  have  been  added  to  the  genus,  including  repre- 
sentatives from  each  of  the  world's  zoogeographic  regions.  However, 
with  the  exception  of  Lee's  (1948)  treatment  of  the  eight  Australasian 
species,  no  comprehensive  paper  on  the  genus  has  yet  appeared. 

A  study  of  the  American  Monohelea  has  resulted  in  the  recognition 
of  7  undescribed  species,  bringing  the  total  for  the  hemisphere  to  13. 
No  doubt  many  more  await  discovery.  These  American  species  fall 
in  four  distinct  groups,  each  with  characteristic  wing  and  leg  markings. 
Within  these  groups  the  species  are  rather  difficult  to  separate,  for  the 
characters  which  prove  useful  in  one  group  may  be  valueless  in  others. 
These  groups  may  also  be  recognized  by  characters  of  the  male  geni- 
talia, which  offer  in  addition  good  specific  characters. 

The  generic  position  of  Ceratopogon  seguax  Williston,  based  on  a 
male  from  St.  Vincent,  cannot  be  determined  with  certainty  until 
the  male  genitalia  or  the  female  are  studied.  According  to  the  original 
description,  seguax  would  probably  fit  as  well  in  Stilobezzia.  Simi- 
larly, the  male  recorded  as  Monohelea  sp.  by  Floch  and  Abonnenc 
(1942)  from  French  Guiana  could  be  either  Monohelea  or  Stilobezzia, 
since  in  their  figure  of  the  male  genitalia  the  aedeagus  is  omitted. 

The  types  of  five  of  the  new  species,  as  well  as  most  of  the  specimens 
studied,  are  in  the  U.  S.  National  Museum.    For  the  loan  or  donation 

'  Bureau  of  Entomology  and  Plant? Quarantine,  Agricultural  Research  Administration,  U.  S.  Depart- 
ment of  Agriculture. 

232992—53  135 


136  PROCEEDINGS    OF    THE    NATIONAL    MUSEUM  ^°^-  "^ 

of  other  specimens  examined  I  am  deeply  grateful  to  Paul  Freeman, 
of  the  British  Museum  (Natural  History),  to  Henry  Dietrich,  of 
Cornell  University,  and  to  John  Lane,  of  the  University  of  Sao  Paulo. 
For  the  generous  gift  or  loan  of  still  other  specimens  I  wish  to  thank 
Mrs.  Elisabeth  C.  Beck  and  J.  A,  Mulrennan  of  Jacksonville,  Florida, 
Bernard  Brookman  of  San  Francisco,  California,  Jean  A.  Laffoon  of 
Ames,  Iowa,  and  Henry  K.  Townes  of  Raleigh,  North  Carolina. 
The  figures  of  the  wings  and  legs  were  made  with  the  aid  of  a  micro- 
projector  and  those  of  the  male  genitalia  with  the  aid  of  an  ocular 
reticule  disc   and  squared  paper.     No   consistent  scale  was  used. 

Family  Heleidae 

Genus  Monohelea  Kieffer,  1917 

Diagnosis. — Body  short,  not  very  haiiy.  Eyes  bare.  Mesonotal 
pits  present  but  small.  Forelegs  and  midlegs  umnodified;  fourth 
tarsal  segments  cylindrical  and  claws  small  and  equal  in  both  sexes; 
hind  leg  with  femur  and  tibia  more  or  less  thickened,  but  without 
spines,  basitarsus  with  a  spine  at  tip,  fourth  segment  cylindrical,  fifth 
segment  in  female  with  one  very  long  claw,  the  other  absent  or  very 
short,  male  claws  small  and  equal  except  hind  claw  as  in  female  in 
tessellata  group;  empodium  absent.  Wings  with  fine  micro trichiae,  a 
few  macrotrichiae  at  wing  tip ;  costa  extending  well  beyond  middle  of 
wing;  two  radial  cells,  second  longer  than  first;  intercalary  fork  fairly 
distinct;  crossvein  r-m  vertical;  median  fork  with  short  stem,  M2 
sometimes  interrupted  at  base;  anal  vein  thickened  in  middle,  but 
without  fold  at  thickening.  Male  genitalia  with  ninth  tergite  tapered, 
the  caudal  margin  truncate  or  broadly  bilobed;  aedeagus  usually  with 
a  pair  of  sharp-pointed,  tapered,  lateral  sclerites  and  an  accessory, 
dorsomedian,  posterior  lobe;  parameres  consisting  of  paired,  sub- 
median  sclerites  of  irregular  shapes. 

Key  to  the  American  species  of  Monohelea 

1.  Wings  without  markings;  mesonotum  shining  black  (subgenus  Schizohelea) . 

1.  leucopeza  (Meigen) 
Wings  with  pronounced  dark  markings;  mesonotum  pruinose,  often  with  mark- 
ings (subgenus  Monohelea) 2 

2.  Hind  femur  and  tibia  yellow,  with  dark  brown  rings  {hieroglyphica  group) .     3 
Hind  femur  and  tibia  shining  black,  only  knee  spot  yellow 9 

3.  Forelegs  and  midlegs  with  distinct  dark  brown  bands;  wings  often  with  exten- 

sive macrotrichiae 4 

Forelegs  and  midlegs  yellowish,  without  distinct  bands;  wings  with  macro- 
trichiae only  at  extreme  distal  margin 6 

4.  Wings  with  macrotrichiae  over  distal  third;  wing  markings  extensive 5 

Wing  without  apparent  macrotrichiae;  wing  markings  reduced,   X-shaped 

marking  in  cells  Mi  and  M2  broken  up  into  3  discrete  spots. 

6.  texana,  new  species 


HELEID    MIDGES,    GENUS   MONOHELEA — ^WIRTH  137 

5.  Wing  with  about  20  small,  scattered,  black  dots  in  addition  to  the  irregular, 

diffuse  gray  markings;  fore  tibia  and  midtibia  dark  only  at  apices;  scutellum 

uniformly  whitish 8.  brasiliensis  Lane 

Wing  with  uniformly  gray,  irregular  markings,  without  small,  black  dots; 
fore  tibia  and  midtibia  entirely  dark;  scutellum  dark  in  middle. 

7.  ornata,  new  species 

6.  Hind  femur  brown  on  basal  third;  scutellum  dark  in  middle;  wing  markings 

not  whitish  bordered 7 

Hind  femur  with  broad  yellow  band  on  sub-basal  fourth;  scutellum  uniformly 
dull  whitish;  wing  markings  bordered  with  whitish  pruinosity. 

3.  maculipennis  (Coquillett) 

7.  Basal  and  median  brown  bands  on  hind  tibia  joined  with  brown,  at  least  on 

ventral  side ;  male  parameres  each  with  2  or  3  distal  lobes 8 

Basal  and  median  brown  tibial  bands  separated  by  a  broad  yellow  band ;  male 
parameres  each  with  simple,  bladelike  apex 5.  macfiei,  new  species 

8.  Sinuate,  dark,  wing  marking  in  cells  Mj  and  M2  entire;  male  parameres  with 

broad,  rounded  distal  lobes 4.  lanei,  new  species 

Sinuate,  dark,  wing  marking  broken  into  3  or  4  separate  spots;  male  parameres 
with  slender,  pointed,  distal  lobes 2.  hieroglyphica  Kieffer 

9.  Scutellum  yellow,  brown  in  middle;  wing  with  2  prominent,  small,  black  spots, 

spot  absent  in  basal  cell;  hind  basitarsus  about  2.5  times  as  long  as  second 
segment ;  mesonotum  yellowish,  with  tiny  brown  dots  (multilineata  group)  10 
Scutellum  entirely  dark  brown  or  yellow  only  in  middle ;  wing  with  two  prom- 
inent anterior  spots  plus  a  spot  in  basal  cell;  hind  basitarsus  1.3  to  2.0  times 
as  long  as  second  segment  {tessellata  group) 12 

10.  Ninth  sternite  of  male  with  4  long  hairs  arising  from  tubercles  in  a  row  at  base 

of  median  lobe 11 

Ninth  sternite  of  male  with  2  long  hairs  arising  from  base  of  median  lobe;  para- 
meres not  connected,  with  irregular,  twisted  apices;  female  abdomen 
pruinose  gray  above,  with  small,  lateral,  shining  brown  spots. 

9.  stonei,  new  species 

11.  Male  aedeagus  with  broad,  bilobed  apex;  parameres  not  connected,  recurved 

apices  long  and  scimitar  shaped;  female  abdomen  pruinose  gray  above,  with 
narrow,  shining  brown  bands  broadening  at  margins  of  segments. 

10.  multilineata  (Lutz) 

Male  aedeagus  cleft  one-third  way  to  base,  with  a  pair  of  long,  slender,  apical 

points;  parameres  broadly  connected  at  bases,  recurved  apices  with  broadly 

rounded  lobes 11.  guianae,  new  species 

12.  Mesonotum  pruinose  gray  with  dark-brown  spots  or  irregular  patches;  scutel- 

lum yellow  in  middle,  with  about  8  marginal  hairs;  wing  bands  distinct  to 

hind  margin,  usually  strongly  interconnected 12.  nebulosa  (Coquillett) 

Mesonotum  dark  brown  with  irregular,  pruinose,  gray  spots;  scutellum  usually 
entirely  black,  with  4  marginal  hairs;  wing  bands  faint  behind  vein  Mj, 
usually  rather  well  separated 13.  johannseni,  new  species 

Subgenus  Schizohelea  Kieffer,  1917 

Schizohelea  Kieffer,  Ann.  Mus  Nat.  Hungarici,  vol.  15,  p.  295,  1917,  vol.  16,  p.  57, 
1918,  and  vol.  17,  p.  89,  1919.— Goetghebuer,  Mem.  Mus.  Hist.  Nat. 
Belgique,  vol.  8,  p.  65,  1920. — Edwards,  Trans.  Ent.  Soc.  London,  vol.  74, 
p.  411,  1926.— Johannsen,  Ann.  Ent.  Soc.  Amer.,  vol.  36,  p.  782,  1943. 
(Genotype,  Cerafopogon  copiosus  Winnertz;  monobasic.) 

Allohelea  Kieffer,  Ann.  Mus.  Nat.  Hungarici,  vol.  15,  p.  364,  1917.  (Genotype, 
Sphaeromyas  pulchripennis  Kieffer;  original  designation.) 


138  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  ^°^-  ^°» 

Following  Goetghebuer  (1934),  I  regard  Schizohelea  as  a  subgenus 
for  leucopeza  Meigen. 

Diagnosis. — This  species  differs  rather  markedly  from  the  other 
species  of  Monohelea  in  having  the  second  branch  of  the  media  broadly 
interrupted  at  the  base,  basitarsus  without  apical  spine,  and  male 
genitalia  with  emarginate  sternite  and  arched  aedeagus. 

1.  Monohelea  (Schizohelea)  leucopeza  (Meigen),  1804 

Figure  18,  m 

Ceratopogon  leucopeza  Meigen,  Klassifikation  und   Beschreibung  .  .  .  Insekten, 

vol.  1,  p.  29,  1804. 
Ceratolophus  leucopeza,  Kieffer,  Genera  insectorum,  fasc.  42,  p.  60,  1906. 
Schizohelea  leucopeza,  Edwards,  Trans.  Ent.  Soc.  London,  vol.  74,  p.  411,  1926. — 

Seguy,  Faune  de  France,  pt.  8,  p.  70,  1937. 
Monohelea    (Schizohelea)    leucopeza,    Goetghebuer,    in  Lindner,  Die  Fliegen  der 

Palaearktischen  Region,  Lief.  78,  p.  53,  1934. 
Ceratopogon  albiiarsis  Wiedemann,  Zool.  Mag.,  vol.  1,  p.  67,  1817. 
Ceratopogon  copiosus  Winnertz,  Linn.  Ert.,  vol.  6,  p.  56,  1852. 
Ceratolophus  copiosus,  Kieflfer,  Genera  insectorum,  fasc.  42,  p.  60,  1906. 
Schizohelea  copiosa,  Kieflfer,  Ann.  Mus.  Nat.  Hungarici,  vol.  15,  p.  295, 1917;  idem, 

vol.  16,  p.  57,  1918;  idem,  vol.  17,  p.  89,  1919.— Goetghebuer,  Mem.  Mus. 

Hist.  Nat.  Belgique,  vol.  8,  p.  66,  1920. — Kieffer,  Faune  de  France,  pt.  11, 

p.  116,  1925. 
Ceratopogon  politus  Coquillett,  Proc.  U.  S.  Nat.  Mus.,  vol.  23,  p.  606,  1901  ( 9 , 

Massachusetts) . 
Ceratolophus  politus,  Kieflfer,  Genera  insectorum,  fasc.  42,  p.  61,  1906. 
Johannseniella  polita,  Malloch,  Bull.  Illinois  State  Lab.  Nat.  Hist.,  vol.  10,  p. 

227,  1914. 
Johannsenomyia  polita,  Malloch,  Bull.  Illinois  State  Lab.  Nat.  Hist.,  vol.  10,  p. 

335,  1915  (New  York). 
Sphaeromias  polita,  Kieflfer,  Ann.  Mus.  Nat.  Hungarici,  vol.  15,  p.  364,  1917. 
Allohelea  polita,  Kieflfer,  Ann.  Mus.  Nat.  Hungarici,  vol.  15,  p.  364,  1917. 
Schizohelea   polita,    Johannsen,   Ann.    Ent.  Soc.   Amer..    vol.  36,  p.   782,   1943 

(?  ^leucopeza  Meigen). 
Bezzia  stecki  Kieflfer,  Broteria,  Ser,  Zool.,  vol.  13,  p.  65,  1915. 

Description. — Female:  Length  1.5  mm.,  wing  1.2  mm.  by  0.5  mm. 
Entirely  shining  black,  only  the  tarsi,  wings,  and  halteres  white. 
Antennae  quite  short,  the  distal  segments  scarcely  elongated. 
Mesonotum  with  scattered,  erect,  long,  black  hairs;  scutellum  with 
six  long  marginal  hairs.  Second  radial  cell  half  again  as  long  as  the 
first,  M2  broadly  interrupted  at  base.  Spermathecae  two,  subequal, 
rather  small  and  subspherical,  the  ducts  sclerotized  a  short  distance. 

Male  (based  on  specimen  from  Suffolk,  England):  Ninth  sternite 
about  three  times  as  broad  as  long,  with  a  shallow,  round  emargination 
in  middle  of  caudal  margin,  not  spiculate;  ninth  tergite  narrow,  sur- 
passing basistyles,  distal  half  with  margins  subparallel,  apex  truncate 
with  setose  apicolateral  corners,  the  inner  surface  coarsely  spiculate. 
Basistyles  broad  at  base,  each  with  a  distinct  lobe  bearing  three  setose 
tubercles  on  inner  side;  abruptly  narrowed  on  distal  half;  dististyles 


HELEID    MIDGES,    GENUS   MONOHELEA — WIRTH  139 

Table  1. — Proportions  of  segments  of  hind  legs  of  female  Monohelea  species 


Species 

Cx 

Tr 

F 

Ti 

Ti 

T2 

T3 

T4 

T5 

1.  leucopeza 

2.  hieroglyphica 

3.  maculipennis 

4.  lanei 

25 
30 
20 
25 

20 
24 
30 

28 
28 
25 

10 
10 
10 
10 
10 
8 
10 
12 
12 
10 

65 
80 
55 
65 
60 
72 
80 
88 
80 
70 

65 
72 
50 
60 
55 
68 
70 
80 
70 
65 

30 
40 
24 
35 
30 
33 
50 
50 
24 
25 

12 
16 
11 
17 
15 
16 
20 
20 
12 
12 

10 

9 

8 

10 

10 

9 

12 

12 

6 

8 

8 
8 
6 
8 
7 
7 
8 
8 
8 
10 

12 
8 
4 
8 

5.  macfiei 

7 

7.  ornata 

9.  stonei 

7 
10 

10.  multilineata 

12.  nebulosa 

13.  johannseni..     

8 

8 

10 

nearly  as  long  as  basistyles,  rather  strongly  curved.  Aedeagus  in 
form  of  a  slender,  rounded  arch  nearly  as  long  as  broad  at  base,  with 
a  pair  of  short,  pointed,  submedian  plates  projecting  dorsocaudad 
at  apex.  Parameres  with  stout,  lateral  apodemes  at  bases;  stems 
stout  and  gently  curved,  contiguous  at  midlength,  their  apices  pointed 
and  abruptly  bent  ventrocephalad  on  about  distal  fourth. 

Type. — Presumably  in  the  Meigen  collection  in  the  Museum 
National  d'Histoire  Naturelle  in  Paris. 

Material  examined. — Massachusetts:  N.  Amherst,  June  1940,  4  99. 

New  York:  Mecklenburg,  June  2,  1940,  A.  Stone,  10  99;  North 
Ridgeway,  June  22,  1940,  S.  C.  Mendall,  1  9. 

Michigan:  St.  Joseph,  May  30,  1938,  C.  W.  Sabrosky,  6  99. 

England:  Corriegills,  Arran,  June  2-4,  1919,  F.  W.  Edwards,  2  99; 
Mildenhall,  Suffolk,  May  22,  1909,  Yerbiu-y,  1  &. 

Remarks. — This  species  is  foimd  in  Europe,  West  Africa,  and 
northeastern  North  America. 

Subgenus  Monohelea  Kieffer,  1917 

Monohelea  Kieffer,  Ann.  Mus.  Nat.  Hungarici,  vol.  15,  p.  295,  1917. — Goetghebuer, 
Mem.  Mus.  Hist.  Nat.  Belgique,  vol.  8,  p.  63,  1920. — Ingram  and  Macfie, 
Ann.  Trop.  Med.  Parasit.,  vol.  15,  p.  344,  1921. — Edwards,  Trans.  Ent.  Soc. 
London,  vol.  74,  p.  410,  1926.— Tokunaga,  Tenthredo,  vol.  3,  p.  156,  1940.— 
Johannsen,  Ann.  Ent.  Soc.  Amer.,  vol.  36,  p.  781,  1943 — Lane,  Arq.  Fac.  Hig. 
Saude  Univ.  Sao  Paulo,  vol.  1,  p.  225,  1948. — Lee,  Proc.  Linn.  Soc.  New 
South  Wales,  vol.  72,  p.  350,  1948.  (Genotype,  Monohelea  hieroglyphica 
Kieffer;  original  designation.) 
Diagnosis. — Wings  with  prominent  markings;  body  with  pruinose 

pattern;  ninth  stemite  of  male  spiculate  and  transverse  or  lobed,  not 

emarginate. 

hieroglyph  ica — group 

Diagnosis. — Hind  legs  conspicuously  banded,  not  markedly  swollen; 
hind  tarsi  slender;  wings  with  extensive  irregular  markings;  male 
aedeagus  broad,  with  median,  anterior  notch  and  triangular,  lateral 
sclerites  with  converging,  sharp-pointed  apices. 


140  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  ^^^-  "^ 

2.  Monohelea  {Monohelea)  hieroglyphica  Kieffer,  1917 

Figure  19,  g 

Monohelea  hieroglyphica  KieflFer,  Ann.  Mus.  Nat.  Hungarici,  vol.  15,  p.  312,  1917 

(cf ,  9  ,  Paraguay). 
Monohelea  hyeroglyphica,  Lane,  Arq.  Fac.  Hig.  Saude  Pub.  Univ.  Sao  Paulo,  vol. 

1,  p.  225,  1948  (Brazil). 

Discussion. — This  is  the  genotype  of  Monohelea.  Macfie's  (1937, 
1940a)  records  of  this  species  from  Trinidad  and  British  Guiana  must, 
I  beUeve,  be  referred  to  macfiei,  new  species,  described  (see.  p.  143), 
from  Louisiana. 

Through  the  kindness  of  John  Lane  I  have  examined  a  female  from 
km.  47,  estrada  Rio-Sao  Paulo,  Rio  de  Janeiro,  Brazil,  collected  in 
February  1945  by  P.  Wygodzinsky.  This  is  from  the  same  locality 
as  the  male  described  and  figured  by  Lane.  Externally  this  species  is 
almost  inseparable  from  M.  (M.)  lanei,  new  species  (see  p.  142),  from 
Florida,  but  the  male  genitalia  are  very  close  to  those  of  M.  (M.) 
maculipennis.  A  sketch  of  the  male  parameres  (after  Lane)  is  included 
here  for  comparison  with  those  of  maculipennis. 

Types. — In  the  Musei  Nationalis  Hungarici  in  Budapest,  cf,  9, 
Paraguay. 

3.  Monohelea  {Monohelea)  maculipennis  (Coquillett),  1905 

Figures  18,  a,  i;  19,  / 

Ceratopogon  maculipennis  Coquillett,  Journ.  New  York  Ent.  Soc,  vol.  13,  p.  64, 
1905  (9  ,  Florida). — Kieffer,  Genera  insectorum,  fasc.  42,  p.  51,  1906. 

Monohelea  maculipennis,  KieflFer,  Ann.  Mus.  Nat.  Hungarici,  vol.  15,  p.  312, 1917. — 
Johannsen,  Ann.  Ent.  Soc.  Amer.  vol.  36,  p.  781,  1943. 

Diagnostic  characters. — A  yellowish  gray,  pruinose  species  with 
faint,  brown,  mesonotal  dots,  pale  j^ellowish  scutellum;  hind  legs  with 
narrow,  dark  rings  and  mngs  with  extensive,  irregular,  grayish  macula- 
tions  bordered  with  whitish  pruinosity. 

Description. — Female:  Length  1  mm.,  wing  1  mm.  by  0.4  mm. 
Head  pale  yellowish,  antennae  and  palpi  bro\^ii ;  proportions  of  flagellar 
segments  of  antennae  12:10:10:12:12:12:12:12:16:16:18:18:24. 
Palpal  segments  in  proportion  of  5 :7 :  10 :5 :  10. 

Mesonotum  grajdsh  pruinose,  with  many  scattered,  small,  brown 
dots  at  the  bases  of  the  dark  mesonotal  haks;  humeri  and  sides  exten- 
sively yellow.  Scutellum  pruinose,  yello-wish  white,  with  four  marginal 
hairs,  the  middle  pair  quite  close  together.  Postscutellum  and  pleura 
pruinose  brown,  the  latter  with  several  indistinct,  transverse,  darker 
lines.  Legs  yellowish,  coxae  and  trochanters  brown,  upper  fourth  of 
midcoxa  and  hind  coxa  yellowish ;  fore  femur  and  midf emur  and  tibiae 
mibanded ;  hind  femur  (figure  18,  i)  brown  at  extreme  base,  two  narrow 
oblique  dark  rings  in  middle  and  a  dark,  preapical,  ventral  spot;  hind 
tibia  mth  narrow  sub-basal,  median,  and  apical  rings;  tarsi  narrowly 


HELEID    MIDGES,    GENUS    MONOHELEA — WIRTH  141 

dark  at  apices  of  segments.    Proportions  of  segments  of  hind  legs  as  in 
table  1. 

Wing  whitish  hyaline,  with  irregular  maciilations,  grayish  by 
transmitted  light,  yellowish  bro^vn  with  narrow,  whitish  pruinose 
borders  by  reflected  light.  Markings  composed  of  a  broad,  irregular 
band  across  wing  at  level  of  first  radial  cell,  filling  basal  half  of  medio- 
cubital  fork,  with  a  prominent,  omega-shaped  spur  in  base  of  cell  Rg; 
a  small  spot  before  middle  of  second  radial  cell;  a  more  or  less  quadrate 
to  X-shaped  mark  across  cell  R5  at  apex  of  second  radial  cell ;  a  sigmoid, 
subapical  mark  across  cells  Mi  and  M2,  often  connected  by  very  narrow 
lines  in  these  cells  to  mesal  band ;  small,  irregular  spots  across  wing  near 
base  and  a  small  spot  past  middle  of  basal  cell.  First  radial  cell  about 
half  as  long  as  second,  a  very  few  macrotrichiae  at  apices  of  cells  R5 
and  Ml,    Halteres  dull  white,  a  black  dot  on  anterior  side. 

Abdomen  dull  whitish,  sides  with  irregular,  dark  patches.  Sper- 
mathecae  two,  very  unequal,  subspherical,  each  with  short  sclerotized 
duct. 

Male:  Ninth  sternite  about  twice  as  broad  as  long,  spiculate, 
posterior  margin  transverse,  abutting  against  base  of  aedeagus,  with 
four  long  hau*s  arising  from  tubercles  in  a  curved  row;  ninth 
tergite  greatly  constricted,  with  sides  subparaUel  on  distal  half,  apex 
truncate,  apicolateral  processes  short.  Basistyles  narrowed  on  distal 
halves ;  dististyles  curved  to  slender,  pointed  apices.  Anterior  margin  of 
aedeagus  broad,  slightly  concave,  with  a  small  median  notch;  aedeagus 
bearing  a  pau-  of  triangular,  submedian  sclerites,  the  bladelike  apices 
of  which  converge  before  level  of  the  irregular  bilobed  apices  of  an 
accessory  pair  of  dorsal  sclerites.  Parameres  with  flaring,  winglike, 
bilobed,  basal  apodemes,  more  or  less  connected  at  midlength  by  a 
pair  of  stout  mesal  lobes,  apices  each  with  a  straight,  long,  slender 
stem  bearing  a  dorsolateral  lobe  about  half  as  long  and  of  same  thick- 
ness, just  beyond  level  of  mesal  bridge. 

Type.—VSlS^M  8366,  9,  Florida,  Jacksonville. 

Material  examined. — Florida:  Crystal  River,  Citrus  County,  July 
17,  1950,  Hudson,  4  99;  Everglades  City,  Collier  County,  Feb.  7, 
1950,  Davidson,  1  cf^,  9  99;  Fort  Myers,  Lee  County,  Feb.  2,  1949, 
Brechtel,  1  cf ;  Islamorada,  Monroe  County,  June  7,  1949,  Smith,  1 
cT,  2  99;  Jacksonville,  date  not  given,  Slosson,  1  9  (type). 

Mexico:  Ciudad  Monte,  Tamaulipas,  Nov.  22,  1943,  Brookman,  1 
&,S  99. 

Guatemala:  Rio  Dulce,  Mar.  21,  1906,  Schwarz  and  Barber,  1  9. 

Panama:  Rio  Trinidad,  June  9,  1902,  Busck,  1  9. 

Remarks. — Formerly  all  the  North  American  Monohelea  with  the 
type  of  wing  pattern  characteristic  of  hieroglyphica  were  called 
maculipennis.     Thus  in  my  (1952)  paper  on  California  Heleidae,  I 


142  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  ^°^-  "^ 

erroneously  included  a  California  record  of  maculipennis  and  also 
cited  several  eastern  records  of  this  species  in  error.  I  have  now 
studied  the  hieroglyphica  group  more  intensively  and  believe  I  have 
correctly  identified  several  component  species  on  the  basis  of  male 
genitalia  supported  by  several  rather  difficult  but  constant  external 
characters  of  the  female.  In  addition  to  hieroglyphica  and  maculi- 
pennis there  appear  to  be  four  or  more  undescribed  species  in  this 
complex.  Although  my  California  specimen  and  that  reported  as 
Monohelea  sp.  by  Johannsen  (1943)  do  not  fit  any  of  those  described 
below,  probably  falling  closest  to  hieroglyphica  in  wing  markings,  a 
positive  diagnosis  of  the  species  must  await  the  collection  of  the  male. 

4.  Monohelea  (Monohelea)  lanei,  new  species 

Figures  18,  b,  j;  19,  e 

Description. — Female:  Length  1.3  mm.,  wing  1.1  mm.  by  0.4  mm. 
As  in  M.  (M.)  maculipennis  (Coquillett),  but  darker  and  slightly 
larger.  Mesonotum  light  brown  pruinose  with  faint  brown  punctures; 
scutellum  brownish  in  middle.  Markings  on  hind  legs  stronger; 
basal  third  of  hind  femur  brown,  a  narrow,  diagonal  band  just  past 
middle,  then  a  small,  brown,  ventral  spot  and  a  strong  preapical  band; 
basal  and  median  bands  on  hind  tibia  broad  and  joined  on  ventral  side, 
the  distal  band  broad.  Proportions  of  segments  of  hind  leg  as  in 
table  1. 

Wing  markings  darker  and  more  extensive  than  in  maculipennis, 
the  omega-shaped  mark  entirely  closed  behind  by  a  broad  extension 
of  the  mesal  band,  the  X-shaped  mark  in  cell  R5  greatly  constricted  in 
middle  and  much  broader  behind.  Macro trichiae  sparse  and  con- 
fined to  wing  margin;  first  radial  cell  about  two-thirds  as  long  as 
second. 

Abdomen  brown  above,  yellowish  on  sides  in  front  and  on  three 
distal  segments.  Spermathecae  two,  large,  very  unequal  and  sub- 
spherical. 

Male:  Ninth  sternite  with  posterior  margin  convex  in  middle, 
spiculate;  ninth  tergite  tapered  to  a  pair  of  blunt,  triangular,  sub- 
median,  caudal  lobes.  Basistyles  narrow,  nearly  straight;  dististyles 
slender,  curved,  with  sharp,  incurved  points.  Aedeagus  with  basal 
halves  broadly  separated  by  a  deep  notch  in  middle  of  anterior  margin, 
lateral  sclerites  with  broad,  triangular  bases,  the  slender,  pointed, 
posterior  portions  directed  caudomesad  and  meeting  each  other  over 
tips  of  a  pair  of  sharp-pointed,  dorsal  processes.  Parameres  a  pair 
of  irregular,  sublateral  sclerites  joined  by  an  indistinct  transverse 
bridge  at  a  third  of  the  distance  from  bases;  basal  apodemes  trilobed, 
directed  laterocephalad  to  bases  of  basistyles;  posterior  portions  with 
subparallel  inner  margins,  tips  each  with  apical,  knoblike,  sclerotized 
lobe  with  slightly  longer,  flattened,  roundly  flaring,  ventral  expansions. 


HELEID    MIDGES,    GENUS    MONOHELEA WIRTH  143 

Types. —Uolotype,  a",  USNM  61091,  Miami,  Florida,  Mar.  25, 
1944,  Wirth.  Allotype  with  same  data  as  type  except  date,  October 
1943.  Paratypes:  Florida:  2  d*  d^,  59  9  ,  same,  except  date,  October 
1943  to  April  1944;  1  9,  Welaka,  June  17,  1946,  Bellamy;  1  9, 
Everglades  City,  Feb.  7,  1950,  Davidson. 

5.  Monohelea  (Monohelea)  macfiei,  new  species 

Figures  18,  c,  h;  19,  h 

Monohelea  hieroglyphica,  Macfie,  not  Kieffer  (misicjentification),  Ann.  Mag.  Nat. 
Hist.,  ser.  10,  vol.  20,  p.  18,  1937  (9  ,  Trinidad);  Ent.  Monthly  Mag.,  vol. 
76;  p.  30,  1940  (cf,  British  Guiana,  genitalia  figured). 

Description.- — Female:  Length  1.2  mm.,  wing  1.4  mm.  by  0.5  mm. 
As  in  M.  (M.)  maculipennis  (Coquillett),  but  the  mesonotum  with  a 
pronounced,  broad,  median,  anterior  band  and  a  large  area  between 
wing  bases  in  front  of  scutellum  uniform  brown,  the  scutellum  with  a 
more  or  less  brownish  median  area.  Abdomen  suffused,  grayish 
brown  above.  Legs  as  figured  by  Macfie  (1937),  brown  bands  on 
hind  legs  quite  distinct;  on  hind  femur  a  band  on  basal  third,  a  diagonal 
band  past  middle,  and  a  narrov/  band  subapically ;  on  hind  tibia, 
narrow  basal,  median,  and  apical  rings;  all  tarsal  segments  with 
narrow  apical  bands. 

Wing  as  in  figure  18,  c,  the  dark  areas  in  the  first  basal  and  anal  cells 
quite  extensive,  the  X-shaped  mark  in  cell  R5  large,  with  the  caudo- 
distal  arm  more  prolonged  than  shown  by  Macfie  and  the  distal  sig- 
moid mark  in  cells  Mi  and  M2  connected  by  broad  lines  in  these  cells 
to  the  broad  median  crossband.  All  wing  markings  brown  by  re- 
flected light,  gray  by  transmitted  light;  a  few  macro trichiae  at  apex 
of  wing. 

Male:  The  resemblance  of  the  Louisiana  male  to  the  male  from 
British  Guiana  figured  by  Macfie  (1940a)  is  remarkable.  The 
slender,  sinuate,  bladelike  parameres  connected  hy  a  narrow  bridge 
at  midlength  are  the  most  distinctive  difi^erence. 

Types. — Holotype,  cf  ,USNM  61092,  allotype,  Kilbourne,  Louisiana, 
May  10,  1947,  Wirth.  Paratypes:  Louisiana:  2  9  9,  with  same  data 
as  type;  1  9,  Baton  Rouge,  May  16,  1947,  Wirth. 

6.  Monohelea  {Monohelea)  texana,  new  species 

Figure  19,  d 

Description. — Male:  Wing  1.1  mm.  long.  Badly  damaged,  anten- 
nae, all  of  legs  but  fore  femur,  midfemm",  and  tibiae  gone.  Mesonotum 
appears  discolored,  all  dark  except  large  quadrate  yellow  patches  on 
humeri;  scutellum  light  yellow,  a  minute  brown  spot  in  middle  of 
anterior  surface.  Forelegs  and  midlegs  yellow,  narrow  basal,  median, 
and  subapical  faint  brown  rings  on  fore  femur;  nan'ow  basal  and 

232992—53 2 


144  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  ^°^-  ^'^ 

broader  median  and  apical  brown  rings  on  fore  tibia;  broad  basal  and 
narrow  median  brown  rings  on  midfemur  and  faint  median  and 
broad,  dark-brown  apical  rings  on  midtibia.  Wings  marked  as  in 
M.  (M.)  hieroglyphica  Kieffer,  the  median  band  broad,  the  omega- 
shaped  spur  distinct  and  open  below,  the  spot  in  cell  Rs  narrow  with 
the  right-hand  arms  of  the  X  absent,  the  sinuate  mark  in  cells 
Ml  and  Mj  reduced  to  three  discrete  spots;  macro trichiae  reduced  to 
a  few  at  v/ing  margin.  Halteres  yellow  with  flat  ends  a.nd  a  spot  on 
anterior  surfaces  of  knobs  dark. 

Ninth  sternite  spiculate,  posterior  margin  broadlj'^  convex  in  middle, 
with  four  long  hairs  arising  from  bases  in  an  arched  row;  ninth  tergite 
tapered,  with  a  pair  of  prominent,  apicolateral  processes.  Aedeagus 
with  basal  halves  narrowly  separated  by  a  deep  notch  in  middle  of 
anterior  margin,  lateral  sclerites  with  broad,  triangular  bases,  the 
slender,  pointed  posterior  apices  meeting  each  other  at  about  level 
of  tip  of  ninth  tergite;  the  dorsal  accessory  structure  biiid  a  third 
the  way  to  base  of  aedeagus,  with  a  pair  of  very  slender  lobes  with 
pointed  apices  curved  ventrad  and  surpassing  apices  of  lateral  sclerites. 
Parameres  joined  together  broadly  near  middle,  each  with  well- 
developed  basal  and  lateral  arms,  stems  abruptly  bent  laterad  two- 
thirds  the  way  to  apices,  then  abruptl}''  bent  ventrad,  with  pointed 
apices  directed  mesocaudad. 

Type. — Holotype,  cf ,  Cornell  Univ.  Type  2787,  Limpia  Canyon, 
Davis  Mountains,  Texas,  July  7,  1917,  Bradley. 

Remarks. — The  male  genitalia  are  most  distinctive,  the  simple 
apices  of  the  parameres  allying  M.  (M.)  texana  with  M.  (M.)  macfiei, 
new  species.  However,  the  external  features,  as  nearly  as  can  be 
ascertained  from  the  badly  damaged  specimen,  are  closer  to  other 
species,  the  wing  markings  being  nearly  like  those  of  M.  (M.)  hiero- 
glyphica Kieffer,  while  the  front  legs  and  midlegs  are  banded  as  in 
M.  (M.)  ornata,  new  species,  and  M.  (M.)  brasiliensis  Lane. 

7.  Monohelea  {Monohelea)  ornata,  new  species 

Figure  18,  d 

Description. — Female:  Length  1.2  mm.,  wing  1.2  mm.  by  0.5  mm. 

Head  pruinose  brown,  vertex  yellowish;  antennae  broken,  pedicel 
brown;  palpi  brown.  Mesonotum  pruinose  gray,  humeri  yellowish; 
a  broad,  median,  longitudinal  band  and  broad  patches  above  wings 
rich,  velvety  brown.  Scutellum  pruinose,  grayish  yellow,  brown  in 
mJddle;  postscutellimi  and  plemm  pruinose  dark  brown.  Fore  femur 
and  midfemur  dull  yellow,  with  faint  median  brown  bands;  fore  tibia 
and  midtibia  brown;  hind  femur  dark  brown  on  basal  third,  a  broad, 
diagonal,  brown  ring  past  middle  and  a  narrow  preapical  brown  ring; 


HELEID    MIDGES,    GENUS   MONOHELEA — WIRTH  145 

hind  tibia  with  broad  basal  and  apical  rings,  and  a  band  just  before 
middle,  brown;  tarsi  dull  yellowish.  Proportions  of  segments  of 
hind  legs  as  in  table  1;  hind  femur  and  tibia  with  rather  long  hairs; 
hind  basitarsus  with  strong  basal  spine.  Claws  on  forelegs  long  and 
equal,  nearly  as  long  as  fom-th  and  fifth  segments  combined;  midtarsi 
broken ;  on  hind  leg  a  single  long  claw  half  again  as  long  as  fifth  segment. 

Wing  grayish  hyaline,  with  very  extensive  gray  maculations  as  in 
figure;  maculations  of  hieroglyphica  type,  but  much  more  extensive, 
with  an  extra  distal,  separate  spot  narrowly  connected  to  the  X-shaped 
mark  in  cell  R5,  and  the  subapical  mark  in  cells  Mi  and  M2  quite  broad. 
First  radial  cell  about  half  as  long  as  second;  macrotrichiae  very  ex- 
tensive, sparsely  covering  distal  third  of  wing  and  including  most  of 
cell  M  3  4.  4  and  anal  cell.  Halteres  not  visible.  Abdomen  uniformly 
dark,  pruinose  brown,  spermathecae  not  examined. 

Types.—Holotjpe,  9,  USNM  61093,  Santa  Rosa  Island,  Escambia 
County,  Florida,  Oct.  10,  1949,  Butler. 

Remarks.-— The  wing,  mesonotal,  and  leg  markings  are  so  distinctive 
that  I  do  not  hesitate  to  describe  M.  (M.)  ornata  from  the  single  female. 

8.  Monohelea  {Monohelea)  brasiliensis  Lane,  1948 

Monohelea  brasiliensis  Lane,  Arq.  Fac.  Hig.  Saude  Pub.  Univ.  Sao  Paulo,  vol.  1, 
p.  226,  1948  (9,  Brazil). 

Description. — ^I  have  not  seen  this  species,  which  is  known  only  from 
the  type  female.  Length  1.2  mm.,  wing  1.3  mm.;  mesonotum  chest- 
nut, yellowish  on  sides  and  in  prescutellar  depression;  scutellum 
whitish.  Legs  yellowish,  fore  femur  dark  at  base,  mudfemur  dark  on 
basal  half;  fore  tibia  and  mid  tibia  dark  at  apices;  hind  femur  with  two 
dark  rings,  one  at  apex  of  basal  third  and  one  in  middle,  hind  tibia  dark 
at  base  and  apex  and  indistinctly  so  in  middle.  Hind  basitarsus  1.8 
times  as  long  as  second  segment.  Wing  with  about  20  black  dots  in 
addition  to  irregular  grayish  markings;  macrotrichiae  numerous  on 
distal  third  of  wing;  first  radial  cell  slightly  over  half  as  long  as 
second  (from  original  description). 

Type. — In  collection  of  University  of  Sao  Paulo,  Brazil,  No.  6781, 
9,  Brazil,  Rio  de  Janeiro,  Estrada  Rio-Sao  Paulo,  km.  47. 

midnlineata — group 

Diagnosis. — Wings  with  two  large  anterior  dark  patches  and  other 
fainter  irregular  markings;  scutellum  dark  in  middle;  legs  dark  with 
yellow  laiee  spots;  hind  legs  slender,  the  tarsi  long  and  unspined; 
ninth  sternite  of  male  with  median,  convex  lobe  and  several  long 
hairs,  aedeagus  long  and  narrow,  parameres  very  long,  with  sharp 
apices. 


146 


PROCEEDINGS    OF    THE    NATIONAL   MUSEUM 


.LANEI 


k.  JOHANNSENI 


W.LEUCOPEZA 


Figure  18. — Species  of  Monohelea:  a-g,  Wings:  a,  maculipennis  (Coquillett);  b,  land,  new 
species;  c,  macfiei,  new  species;  d,  ornata,  new  species;  e,  stonei,  new  species;  /,  nebulosa 
(Coquillett);  g,  johannseni,  new  species,  h-k,  Hind  legs  of  females:  h,  macfiei,  new  species; 
i,  maculipennis  (Coquillett);  ;',  lanei,  new  species;  k,  johannseni,  new  species.  /,  Sperma- 
thecae,  johannseni,  new  species,     m,  Male  genitalia,  leucopeza  (Meigen). 


HELEID    MIDGES,    GENUS    MONOHELEA — ^WIRTH  147 


STONEI 


J.  MULTILINE  ATA 


*.  GUIANAE 


Figure  19. — Species  of  Monohelea,  male  genitalia;  a,  johannseni,  new  species;  h,  tessellata 
(Zetterstedt);  c,  nehulosa  (Coquillett);  d,  texana,  new  species;  e,  lanei,  new  species;  /, 
maculipewiis  (Coquillett);  g,  hieroglyphica  Kieffer;  male  parameres  (after  Lane);  h,  macfiei, 
new  species;  t,  stonei,  new  species;  ;,  multilineata  (Lutz);  k,  guianae,  new  species. 


148  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  ^°^-  "^ 

9.  Monohelea  {Monohelea)  stonei,  new  species 

Figures  18,  e;  19,  i 

Diagnostic  characters. — A  yellowish  gray,  pruinose  species  with 
maculate  wings;  scutellum  yellow  with  brown  center;  legs  dark, 
abdomen  dull  gray  pruinose  above. 

Description. — Female:  Length  2.0  mm.,  wing  1.5  mm.  by  0.7  mm. 
Head  yellowish,  vertex  pruinose,  pale  gray;  antenna  with  pedicel  and 
bases  of  flagellar  segments  yellow,  apices  of  short  segments  and  all  of 
long  distal  segments  and  palpi  brown;  proportions  of  flagellar  seg- 
ments 15:10:10:10:10:10:12:12:18:18:18:18:24.  Palpal  segments 
in  proportion  of  5 :8 :  15 :  10 :  15,  third  segment  scarcely  swollen,  with  a 
rather  large  sensory  pit  at  midlength. 

Mesonotum  yellowish  gray  pruinose,  with  tiny  brown  dots  at  the 
bases  of  the  brownish  mesonotal  hairs.  Scutellum  yellow,  brown  in 
middle,  with  four  moderate  marginal  hairs  and  a  few  fine  setae. 
Postscutellum  and  pleura  pruinose  dark  brown.  Coxae  brown; 
trochanters,  femora,  and  tibiae  light  brown  on  forelegs  and  midlegs, 
black  on  hind  legs;  all  knees  prominently  yellowish,  tarsi  yellow. 
Hind  femur  and  hind  tibia  moderately  thickened;  basitarsus  with  a 
slender  spine  at  base,  distal  tarsal  segments  on  hind  legs  each  with 
a  fine  apical  spine;  a  single  long  claw  on  each  hind  leg.  Proportions 
of  segments  of  hind  leg  as  in  table  1 . 

Wing  grayish  hyaline,  with  two  prominent,  black  spots  and  fainter 
posterior  infumation  as  figured;  fu-st  radial  cell  about  half  as  long  as 
second.     Halteres  infuscated. 

Abdomen  dark  brown,  entire  dorsum  pruinose  gray  except  for  a 
pair  of  small  lateral  spots  and  a  pair  of  submedian  sensory  dots  on 
each  segment,  polished  brown.  Spermathecae  two,  subequal,  ovoid, 
each  with  a  very  short  sclerotized  duct. 

Male:  Similar  to  the  female,  with  the  usual  sexual  differences; 
antennal  plume  golden  at  base,  dark  brown  toward  apex.  Ninth 
sternite  three  times  as  broad  as  long,  with  a  narrow,  median,  poste- 
rior lobe  bearing  two  long  hau-s  extending  into  the  concave  base  of  the 
aedeagus;  sternite  spiculate  on  posterior  half;  ninth  tergite  as  long  as 
basistyles,  tapered  to  the  truncated  apex,  bearing  a  pair  of  short, 
rounded,  apicolateral  processes.  Basistyles  about  twice  as  long  as 
broad,  simple,  very  convex  on  outer  sides;  dististyles  about  0.7  as  long 
as  basistyles,  nearly  straight  and  stout  to  apices.  Aedeagus  with 
base  as  broad  as  the  lobe  of  ninth  sternite,  a  slight  sub-basal  swelling, 
a  second  swelling  at  midlength,  with  distal  half  cleft  into  a  pair  of 
attenuated,  submedian,  pointed  blades  attaining  0.8  length  of  basi- 
styles.    Parameres  consisting  of  a  pau'  of  very  irregular  processes. 


HELEID    MIDGES,    GENUS    MONOHELEA — ^WIRTH  149 

bases  of  each  with  a  large  anterior  and  lateral  apodeme,  stem  rather 
stout  and  crooked,  distal  portion  abruptly  bent  ventrocephalad,  then 
greatly  attenuated  and  bent  ventrocaudad  to  a  sharp  point. 

Types. — Holotype,  9,  USNM  61094,  Baton  Rouge,  Louisiana,  May 
16,  1947,  Wirth.  Allotype  with  same  data  as  type  except  date. 
May  3,  1947.  Paratypes:  Louisiana:  ecfcf ,  3  99,  with  same  data  as 
t3npe  except  date,  May  6-20,  1947.  Mississippi:  4  cT"  d",  10  99,  Horn 
Island,  June  and  July  1944,  E.  A.  Richmond.  Florida:  11  99,  Gray- 
ton  Beach,  Walton  County,  May,  Sept.,  1949,  Butler;  1  9,  Santa 
Rosa,  Walton  County,  July  19,  1949,  Peterson;  8  cf  cT,  21  99,  In- 
nerarity  Point,  Escambia  County,  May,  1950,  Rathert;  8  99,  Santa 
Rosa  Island,  Escambia  County,  July  7,  1949,  Butler;  5  99,  Panama 
City  Beach,  Bay  County,  May  6,  1949,  McElvey;  5  99,  Crystal  River, 
Citrus  County,  July  17,  1950,  Hudson;  3  99,  Fort  Myers,  Lee  County, 
Oct.  8,  1948,  Brechtel;  3  99,  Everglades  City,  May,  Oct.,  1948,  1950, 
Huntoon;  1  cf ,  1  9,  Big  Pine  Key,  Monroe  County,  Apr.  1,  1950, 
Sermon.  Georgia:  2  cf  cf,  4  99,  Thomasville,  May  15-30,  1949, 
Palmer.  Tennessee:  1  9,  Nashville,  June  18,  1937,  Adams.  Iowa: 
1  9,  Sioux  City,  July  15,  1950,  Laffoon.  Virginia:  3  d"  cf^,  6  99,  Falls 
Church,  July  8,  29,  1950,  Wirth.  Maryland:  4  99,  Leeds,  Dorchester 
County,  July  10,  1907,  Barber.  New  Jersey:  1  9,  Newport,  July  1, 
1937  (light  trap).  Costa  Rica:  1  cf ,  Higuito,  San  Mateo,  no  date, 
Schild.  Panamd:  1  9,  Fort  Kobbe,  Canal  Zone,  Aug.  21,  1950, 
Carpenter. 

10.  Monohelea  (Monohelea)  multilineata  (Liitz),  1914 

Figure  19,  j 

Palpomyia  multilineata  Lutz,  Mem.  Inst.  Oswaldo  Cruz,  vol.  6,  p.  93,  1914  (Brazil). 
Monohelea  multilineata,  Johannsen,  Ann.  Ent.  Soc.  Amer.,  vol.  36,  p.  781,  1943. — 
Lane,  Rev.  Ent.,  vol.  16,  p.  368,  1945. 

Lane  (1945)  has  very  adequately  redescribed  this  Brazilian  species. 
It  resembles  M.  (M.)  stonei,  new  species,  in  general  coloration  and 
structure,  the  mesonotum  yellowish  with  brown  dots,  the  scutellum 
yellow  with  median  brown  spot  and  four  marginal  bristles,  forelegs 
and  midlegs  yellowish,  the  hind  femur  and  tibia  black  and  scarcely 
swoUen  and  hind  tarsi  unspined.  The  wings,  however,  in  addition 
to  the  two  anterior  black  spots,  have  smaller,  rather  strong  spots  at 
the  wing  margin  m  cells  R5  and  Mi,  and  across  the  apex  of  cell  M2. 
The  polished,  brown,  lateral  spots  on  the  abdomen,  which  are  small 
in  stonei,  are  larger  and  narrowly  connected  across  the  anterior  margin 
of  each  tergite  in  multilineata. 

The  male  genitalia  of  multilineata  are  of  the  same  type  as  those  of 
stonei,  but  there  are  four  rather  than  two  bristles  at  the  base  of  the 


150  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  ^®^-  ^<" 

lobe  of  the  ninth  sternite,  the  aedeagus  is  broader  at  the  apex  and 
only  slightly  notched  rather  than  deeply  cleft,  and  the  parameres  are 
slender  and  nearly  straight  with  the  apices  abruptly  bent  and  scim- 
itar-shaped. 

Material  examined. — Brazil :  Km.  47,  Estrada  Rio-Sao  Paulo,  Rio  de 
Janeiro,  Brazil,  February  1945,  Wygodzinsky,  2  cfcf ,  2  99. 

Remarks. — Monohelea  nigeriae  Ingram  and  Maciie,  1922,  known 
from  the  female  from  West  Africa,  is  closely  related  to  M.  (M.)  stonei 
and  to  M.  (M.)  muUilineata,  but  according  to  the  original  description, 
the  tarsi  bear  ventral  spines,  as  in  the  tessellata  group,  and  the  sperma- 
thecae  are  imequal  in  size. 

11.  Monohelea  (Monohelea)  guianaef  new  species 

Figure  19,  A; 

Monohelea  muUilineata,   Macfie,  not  Lutz   (misidentification) ,   Proc.   Ent.  Soc. 
London,  Ser.  B,  vol.  9,  p.  187,  1940  (c?,  British  Guiana). 

Macfie  (1940)  has  given  as  satisfactory  a  description  of  this  insect 
as  was  possible  from  the  single  male  specimen  available.  This  speci- 
men was  borrowed  from  the  British  Museum  for  study  and  externally 
cannot  be  separated  from  either  M.  (M.)  muUilineata  (Lutz)  or 
M.  (M.)  stonei,  new  species.  However  the  male  genitalia  are  quite 
distinct  and  may  be  characterized  as  follows:  Ninth  sternite  very 
short,  with  a  low  median  lobe  on  caudal  margin  fitting  against  base  of 
aedeagus,  spiculose,  with  four  long  hairs  arising  from  tubercles  in  a 
line  at  base  of  median  lobe;  nmth  sternite  very  narrow,  tapered  to 
tip,  with  a  pair  of  short,  nipplelike  apieolateral  processes.  Basistyles 
about  twice  as  long  as  broad;  dististyles  nearly  straight,  with  blimt, 
scarcely  narrowed  apices.  Aedeagus  very  narrow,  about  2.5  times  as 
long  as  broad  at  base,  the  anterior  arch  very  low,  apex  cleft  about  a 
third  way  to  base,  with  a  pair  of  very  slender,  submedian  processes, 
from  the  bases  of  which  a  pair  of  slender,  lateral  arms  curve  around 
stems  of  parameres.  Parameres  broadly  joined  together  at  bases, 
each  with  a  short  anterior  and  lateral  arm,  stems  slender,  apex  of  each 
thickened  and  abruptly  bent  laterocephalad  in  a  broadly  rounded 
lobe  about  half  as  long  as  dististyle. 

Types. — Holotype,  cf,  British  Museum  (Natural  History)  collec- 
tion, Mazaruni,  British  Guiana,  Aug.  21,  1937,  second  growth  (low 
forest),  Richards  and  Smart. 

tessellata — group 

Wings  with  three  large  dark  anterior  spots  and  irregular  posterior 
infuscated  areas;  scutellum  often  with  middle  yellow;  hind  legs  swollen 
and  black,  with  yellow  knees,  the  tarsi  short  and  with  strong  ventral 


HELEID    MIDGES,    GENUS    MONOHELEA — WIRTH  151 

spines;  claw  of  hind  leg  of  male  long  and  single  as  in  the  female;  male 
aedeagus  with  slender,  pointed,  posterior  sclerite  borne  in  a  notch  in 
the  quadrate  basal  sclerite,  parameres  short,  curved,  and  pointed. 

12.  Monohelea  {Monohelea)  nebulosa  (Coquillett),  1901 

Figures  18,/;  19,  c 

Ceratopogon  nebulosus  Coquillett,  Proc.  U.  S.  Nat.  Mus.,  vol.  23,  p.  606,  1901 

(cf,  New  Jersey). 
Ceratolophus  nebulosus,  Kieffer,  Genera  insectorum,  fasc.  42,  p.  60,  1906. 
Johannseniella  nebulosa,  Malloch,  Bull.  Illinois  State  Lab.  Nat.  Hist.,  vol.  10, 

p.  226,  1914. 
Hartornyia  nebulosa,  Malloch,  Bull.  Illinois  State  Lab.  Nat.  Hist.,  vol.  10,  p.  340, 

1915  (Indiana). 
Monohelea  nebulosa,  KieflFer,  Ann.  Mus.  Nat.  Hungarici,  vol.  15,  p.  312,  1917. — 

Johannsen,   Ann.   Ent.   Soc.  Amer.,  vol.  36,  p.  781,  1943   (Massachusetts, 

Idaho) . 

Description. — Female:  Length  2.1  mm.,  wing  1.7  mm.  by  0.6  mm. 
Head  dull,  dark  brown,  with  a  few  pruinose  gray  spots  on  vertex; 
antennae  light  brown,  distal  segments  not  darker.     Palpi  light  brown. 

Mesonotum  pruinose  gray,  with  scattered,  small,  dark-brown  dots, 
each  mesonotal  hair  arising  from  a  dot,  these  dots  more  or  less  con- 
fluent, especially  on  posterior  portion  of  mesonotum.  Scutellum 
brown,  middle  third  yellow,  with  about  eight  marginal  bristles. 
Postscutellum  and  plem-a  dark  brown.  Legs  shining  dark  brown, 
knees  and  tarsi  yellowish.  Posterior  femora  and  tibiae  greatly 
thickened;  hind  basitarsus  with  basal  and  distal  spine;  second  segment 
with  two  distal  spines,  third  segment  with  one  distal  spine;  claws 
minute  and  equal  on  forelegs  and  midlegs,  outer  claw  very  long, 
four  times  the  length  of  inner,  on  hind  legs.  Proportions  of  segments 
of  hind  leg  as  in  table  1. 

Wing  grayish  hyaline,  a  quadrate  black  patch  across  first  basal  cell 
at  half  its  length;  two  broad,  irregular,  more  or  less  interconnected, 
transverse  bands  across  wing,  the  first  at  first  radial  cell,  the  second  at 
apex  of  second ;  cell  Rs  thus  with  a  dark,  a  light,  a  dark  and  a  light  band, 
all  of  subequal  breadth.  First  radial  cell  0.4  times  as  long  as  second. 
Halteres  pale  yellow. 

Abdomen  uniform,  pruinose,  dark  brown;  anal  segment  yellow. 

Male:  Ninth  sternite  transverse,  without  caudal  lobe,  posterior 
portion  spiculate;  ninth  tergite  very  narrow,  with  sides  subparallel  on 
distal  half,  apicolateral  processes  short  and  rounded.  Basistyles  very 
stout  and  convex;  dististyles  slender  and  curved,  0.6  as  long  as  basi- 
styles. Aedeagus  with  two  sections,  basal  part  about  as  broad  as  long, 
anterior  margin  contiguous  with  sternite,  posterior  margin  deeply 
emarginate,  the  slender,  rodlike,  distal  sclerite  with  notched  apex  fit- 


152  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  ^°^-  ^°^ 

ting  into  the  emargination.  Parameres  connected  by  a  transverse 
bridge  near  bases,  with  slender  anterior  and  lateral  apodemes;  each 
paramere  with  posterior  portion  narrowed  and  curved  ventrolaterad 
to  a  bladelike  apex,  a  small  sub-basal  tooth  on  ventral  side. 

Material  examined.— New  York:  Farmingdale,  July  9,  1938,  H.  and 
M.  Townes,  1  cf ;  Millwood,  June  21,  1936,  H.  Townes,  1  9. 

New  Jersey:  Atsion,  June  17,  1949,  H.  Townes,  1  9;  Medford  Lakes, 
June  17,  1939,  H.  Townes,  1  cf;  Riverside,  June  18,  1939,  H.  Townes, 
1  9;  Riverton,  June  19,  Johnson,  1  cf  (type). 

Georgia:  Thomasville,  May  15-30,  Pahner,  1  9. 

Arkansas:  Pike  County,  June  12,  1938,  Turner,  1  9. 

Remarks. — This  species  is  very  similar  to  the  Palearctic  species  M. 
(M.)  tessellata  (Zetterstedt),  1850,  which,  however,  has  on  the  scutel- 
lum  four  marginal  bristles,  the  ends  of  which  are  yellowish,  as  is  the 
median  area.  The  wing  markings  of  tessellata  are  much  more  nearly  as 
in  M.  (M.)  johannseni,  new  species. 

A  male  of  tessellata  from  Norfolk,  Hatfield,  England,  was  examined 
through  the  kindness  of  Paul  Freeman  of  the  British  Museum  (Natural 
History) .  The  specimen  is  unsatisfactory  for  a  description  of  external 
characters,  as  it  was  glued  to  a  card  when  wet.  It  is  noted,  however, 
that  both  the  first  and  second  segments  of  the  hind  tarsi  bear  a  distal 
pair  of  spines.  The  aedeagus  of  the  genitalia  consists  of  a  heavily 
sclerotized  basal  plate  about  three  times  as  broad  as  long,  bearing  a 
small,  crescentic,  median  sclerite  on  the  ventroposterior  margin  and 
bearing  dorsoposteriorly  a  large,  triangular  structure  with  sharp- 
pointed,  median  apex,  and  widely  flaring,  lateral  arms  articulating  with 
inner  margins  of  basistyles.  Parameres  with  slender  anterior  arms 
connected  by  a  broad,  median  bridge,  the  posterior  portions  simple, 
the  basal  halves  stout,  the  distal  halves  very  slender  and  curved  ventro- 
cephalad. 

According  to  Tokunaga's  (1940)  description,  his  record  of  tessellata 
from  Japan  probably  refers  to  an  as  yet  unnamed  species.  This  Jap- 
anese species  has  the  scutellum  yellow  only  in  the  middle,  as  in  nebulosa, 
which  it  also  closely  resembles  in  wing  markings;  but  the  mesonotura 
has  distinct  yellow  spots,  the  antennae  have  the  distal  segments  and 
the  apices  of  the  basal  segments  brownish,  the  legs  are  yellowish  brown 
with  dark  brown  markings,  and  the  dorsum  of  the  abdomen  is  pale, 
yellowish  brown. 

The  West  African  species  M.  (M.)  litoraurea  Ingram  and  Macfie, 
1921,  (female)  and  M.  (M.)  mimas  de  Meillon,  1939,  (male)  are  very 
similar  to  M.  (M.)  nebulosa  (Coquillett) .  The  genitalia  of  mimas  are 
the  same  as  those  of  nebulosa  except  that  the  dististyles  are  abruptly 
bent  near  the  base. 


HELEID    MIDGES,    GENUS    MONOHELEA — WIRTH  153 

13.  Monohelea  {Monohelea)  johannseni,  new  species 

Figures  18,  g,  k,  I;  19,  a 

Monohelea  tessellata,  Johannsen,  not  Zetterstedt   (misidentification) ,  Ann.  Ent. 
Soc.  Amer.,  vol.  36,  p.  781,  1943  (Alabama). 

Diagnostic  characters. — A  dark  brown  species  with  large  pruinose 
gray  mesonotal  markings,  black  scutellum  and  legs,  and  maculate 

WIQgS. 

Description. — Female:  Length  1.5  mm.,  wing  1.2  mm.  by  0.5  mm. 
Closely  resembling  M.  (M.)  nebulosa  (Coquillett)  but  smaller,  the 
mesonotum  dark  brown  with  large,  irregular,  pruinose  gray  areas,  the 
scutellum  entirely  blackish  and  with  four  long  and  several  shorter 
marginal  hairs.  Proportions  of  hind  leg  of  female  as  in  table  1,  hmd 
tarsi  with  one  basal  and  two  distal  spines  on  fu'st  segment,  two  spines 
at  tip  of  second  and  occassionally  two  spines  at  apices  of  third  and 
fourth  segments;  inner  claw  on  hind  leg  of  female  about  a  thu'd  as 
long  as  outer.  Wing  as  in  nebulosa,  but  the  markings  behind  vein 
iVli  much  fainter  than  those  on  anterior  part  of  wing.  Spermathecae 
two,  subequal,  oval,  each  with  very  short,  sclerotized  duct. 

Male  genitalia  as  in  nebulosa,  but  the  parameres  simple,  without 
the  sub-basal  ventral  tooth,  and  distinctly  and  abruptly  bent  mesad 
at  distal  fifth. 

Types.— Roloiype,  9,  USNM  61095,  Falls  Church,  Vkginia,  July  8, 
1950,  Wirth.  Allotype,  Innerarity  Point,  Escambia  County,  Florida, 
Apr.  29,  1949,  Kathert.  Paratypes:  Virginia:  2  99,  Mountain  Lake, 
July  15,  1938,  July  21,  1940,  L.  and  M.  Mihie.  Florida:  4  cf  cf ,  5  99, 
same  data  as  allotype  except  date  April  and  May  1949;  1  9,  Santa 
Rosa,  Walton  County,  May  5,  1950,  Peterson.  Alabama:  1  9,  La- 
Place,  June  9,  1917,  Bradley.  Michigan:  1  9,  Cheboygan  County, 
July  17,  1942,  Sabrosky.  Iowa:  1  9,  Pikes  Peak  State  Park,  Clayton 
County,  July  4,  1949,  Laffoon. 


References 

Floch,  H,,  and  Abonnenc,  E. 

1942.  Ceratopogonides  divers  de  la  Guyane  Francaise  III,     Inst.  Pasteur 

Guyane  Terr.  L'Inini  Publ.  55,  6  pp.,  2  figs. 

GOETGHEBUER,  M. 

1934.    Heleidae  (Ceratopogonidae),  In  Lindner,  Die  Fliegen  der  Palaearkti- 
schen  Region,  Lief.  78,  pp.  49-94. 

JOHANNSEN,  O.  A. 

1943.  A  generic  synopsis  of  the  Ceratopogonidae  (Heleidae)  of  the  Anaericas, 

a  bibliography,  and  a  list  of  the  North  American  species.     Ann. 
Ent.  Soc.  Amer.,  vol.  36;  pp.  763-791. 

KlEFFER,  J.  J. 

1917.    Chironomides  d'Amerique  conserves  au  Musee  National  Hongrois  de 
Budapest.     Ann.  Mus.  Nat.  Hungarici,  vol.  15,  pp.  292-364,  43  figs. 
Lane,  J. 

1945.    Redescrigao  de  Ceratopogonideos  Neotr6picos  (Diptera:  Ceratopogo- 
nidae).    Rev.  Ent.,  vol.  16,  pp.  357-372,  19  figs. 
Lee,  D.  J. 

1948.    Australasian  Ceratopogonidae  (Diptera,  Nematocera).     Part  4.     The 
Stilobezzia  group  of  genera.     Proc.  Linn.  Soc.  New  South  Wales, 
vol.  72,  pp.  345-356,  1  pi.,  23  figs. 
Macfie,  J.  W.  S. 

1937.    Ceratopogonidae  from  Trinidad.     Ann.  Mag.  Nat.  Hist.,  ser.  10,  vol. 

20,  pp.  1-18,  6  figs. 
1940a.    A  report  on  a  collection  of  Ceratopogonidae  (Diptera)  from  British 

Guiana.     Ent.  Monthly  Mag.,  vol.  76,  pp.  23-32,  4  figs. 
1940b.    Ceratopogonidae    (Diptera)    from    British    Guiana    and    Trinidad. 
Proc.  Ent.  Soc.  London,  ser.  B,  vol.  9,  pp.  179-195,  4  figs. 
Tokunaga,  M. 

1940.    Chironomoidea  from  Japan    (Diptera),   XII.     New  or  little-known 
Ceratopogonidae  and  Chironomidae.     Philippine  Journ.  Sci.,  vol. 
72,  pp.  255-311,  4  pi. 
WiRTH,  W.  W. 

1952.    The  Heleidae  of  California.     Univ.  California  Publ.  Ent.,  vol.  9,  pp. 

95-266,  33  figs. 
154 


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PROCEEDINGS   OF   THE   UNITED  STATES   NATIONAL  MUSEUM 


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U.  S.  NATIONAL  MUSEUM 


Vol.103  Washington  :  1953  No.  3321 


A  REVIEW  OF  THE  BEETLE  FAMILY  CEPHALOIDAE 


By  Ross  H.  Arnett,  Jr.^ 


The  family  Cephaloidae  consists  of  only  one  genus,  Cephaloon 
Newman,  1838,  with  eight  known  species  that  are  locally  rather  com- 
mon but  are  known  only  from  the  Eastern  United  States,  Western 
United  States,  Western  Canada,  Japan,  and  Amur.  All  of  the  species 
are  closely  related  and  quite  variable  in  color.  Useful  separation 
characters  are  to  be  found  in  the  antennae  and  various  male  structures. 
As  yet  Httle  is  known  concerning  the  habits  and  life  histories  of  mem- 
bers of  this  group.  All  known  species  of  this  family  are  represented 
in  the  collections  of  the  U.  S.  National  Museum. 

After  study  of  some  material  from  the  Ussuri  River  Valley  recently 
sent  to  me  for  determination,  it  seems  desirable  to  reconsider  the 
taxonomy,  affinities,  and  distribution  of  this  group.  The  ranking  of 
this  small  assemblage  of  species  as  a  family  on  equal  ground  with  the 
rest  of  the  families  in  the  order  is  still  somewhat  doubtful  in  my  mind. 
I  believe  the  placing  of  the  Cephaloidae  as  a  satellite  of  the  great 
family  Tenebrionidae  is  firmly  established,  but  my  studies  in  the 
Tenebrionoidea  have  not  as  yet  revealed,  on  the  basis  of  our  present 
ranking  and  evaluation  of  what  defines  a  family,  any  group  with  which 
the  genus  Cephaloon  can  be  incorporated;  hence,  I  retain  it  as  a  family. 

I  wish  to  thank  Mr.  Hugh  B.  Leech  and  Dr.  E.  C.  Van  Dyke,  both 
of  the  California  Academy  of  Sciences,  and  Dr.  Hans  Klapperich  of 
Bonn,  Germany,  for  the  loan  of  several  specimens  used  in  this  study. 

'  Bureau  of  Entomology  and  Plant  Quarantine:  U.  S.  Department  of  Agriculture. 

232728—63  155 


156  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  io3 

Family  Cephaloidae  LeConte 

Cephaloidae  LeConte,  Smithsonian  Misc.  Coll.,  vol.  3,  art.  3,  Classification  of  the 
Coleoptera  of  North  America,  pt.  1,  p.  259,  1862. 

LeConte  originally  proposed  this  family  for  the  North  American 
Cephaloon  lepturides  Newman,  but  also  referred  to  two  species,  al- 
though not  by  name,  which  had  been  described  by  Motschulsky  from 
the  Amur  River  Valley.  Several  genera  have  since  been  erected  for 
species  in  this  group,  and  then  synonymized,  until  in  the  present 
study  the  family  contains  only  the  original  genus  Cephaloon  Newman. 

Family  diagnosis. — Size  8-20  mm. ;  head  elongate,  diamond-shaped, 
deflexed;  antennae  11-segmented,  filiform,  with  apical  segments  some- 
what enlarged,  inserted  between  the  eyes  and  bases  of  the  mandibles; 
mandibles  elongate,  acute  at  apex,  never  bifid  or  otherwise  modified; 
eyes  reniform,  not  prominent;  maxillary  palpus  4-segmented,  first 
segment  small,  obscure,  apical  segment  triangular.  Pronotum  with- 
out lateral  margins,  smooth,  always  abruptly  narrowed  anteriorly 
from  the  middle.  Legs  slender;  front  coxal  cavities  open  behind; 
front  and  middle  coxae  prominent,  conical;  apical  spurs  of  all  tibiae 
large,  two  on  each  tibia;  tarsi  5-5-4,  the  segments  all  simple,  not  lobed 
or  tomentose  beneath;  claws  pectinate,  with  a  subequal  membranous 
lobe  beneath  each  claw.  Elytra  with  vague  costae,  minutely  punctate, 
never  striate.  Abdomen  with  5  visible  sternites  in  the  female,  7  in 
the  male  (sternites  2  +3  to  9  visible  in  the  male).  Body  covered  with 
very  fine  pubescence. 

Male  genitalia:  Apical  abdominal  segments  (7-9)  of  the  male 
considerably  modified.  Segment  7  with  sternite  and  tergite  laterally 
fused,  forming  a  globular  shaped  segment.  Segment  8  with  sternite 
and  tergite  laterally  fused,  forming  two  laterally  triangular  pieces. 
Segment  9  closely  fitted  into  segment  8,  the  tergite  and  sternite  both 
triangular  and  filling  the  emarginations  of  segment  8.  Genital  organs 
themselves  quite  simple;  paramere  two  short,  freely  articulate  lobes 
fitted  onto  the  apex  of  the  large  troughlike  basal  piece  which  more  or 
less  envelops  a  simple,  small,  somewhat  curved,  and  tubelike  median 
lobe;  no  evidence  of  a  tegminite. 

Affinities  of  the  family. — The  heteromerous  tarsi  place  this  family 
in  the  Tenebrionoidea;  the  open  anterior  coxal  cavities  place  itjn^the 
group  including  the  Oedemeridae,  Pythidae,  and  Serropalpidae,  as 
distinguished  from  the  group  including  the  Alleculidae,\Lagriidae, 
and  Tenebrionidae,  all  of  which  have  closed  coxal  cavities.  The  ser- 
rate claws  with  fleshy  pulvilli  and  the  laterally  fused  eighth  abdominal 
segment  of  the  male  separate  it  from  the  others  of  this  group  of  Teneb- 
rionoidea. The  very  small  median  lobe  and  the  larger  paramere 
with  the  small  lateral  lobes  show  affinities  with  the  genus  Mycterus  of 


BEETLE  FAMILY  CEPHALOIDAE — ARNETT 


157 


U 


(^ 


158  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  103 

the  family  Pythidae.  In  addition,  Mycterus  and  Cephaloon  both 
have  prominent  procoxae  and  mesocoxae  and  the  prothorax  is  with- 
out lateral  margins.  Most  workers  in  the  past  have  considered  the 
Oedemeridae  and  Cephaloidae  closely  allied,  but  my  incomplete 
studies  of  the  male  genitalia  of  the  heteromerous  beetles  indicate  that 
this  is  not  so.  The  very  different  structure  of  the  apical  abdominal 
segments  of  the  species  of  Cephaloon  leads  me  to  suppose  that  these 
species  are  far  removed  from  the  oedemerids,  but  their  true  affinities 
are  not  yet  recognized. 

Notes  on  the  distribution  oj  the  species. — As  can  be  seen  from  the 
accompanying  map  (fig.  20),  the  species  of  this  family  show  the  typical 
distribution  pattern  of  a  Holarctic  group  which  was  probably  derived 
from  some  Asian  stock  and  spread  to  North  America  via  the  Alaskan 
land  bridge.  It  is  interesting  to  note  that,  based  on  both  external 
morphology  and  the  morphology  of  the  male  genitalia  (in  the  case 
of  C.  pollens)  both  of  the  Asian  species  are  most  similar  to  two  of  the 
Eastern  North  American  species  and  not  to  the  Western  North 
American  species.  If  these  similarities  of  morphology  reflect  relation- 
ship, as  wo  assume  they  do,  then  we  again  see  the  often-repeated 
pattern  of  a  pre-glacial  distribution  across  northern  Canada  from 
Alaska,  down  through  Ontario  and  into  Eastern  United  States. 
This  I  believe  to  be  a  further  stock-piling  of  evidence  against  the 
theories  of  continental  drift  and  the  North  Atlantic  land  bridge  idea. 

Genus  Cephaloon  Newman 

Ichnodes  Dejean,  1834,  Cat.  Col.  ed.  3,  p.  227.  {Nomen  nudum,  one  ms.  trivial 
name  listed.) 

Cephaloon  Newman,  Ent.  Mag.,  vol.  4,  p.  376,  1838.  (Genotype,  Cephaloon 
lepturides  Newman;  1838;  monobasic.) 

Cephalaon  Motschulsky  in  Schrenck,  Reisen  und  Forschungen  in  Amurlande, 
vol.  2,  pt.  2,  p.  140,  I860.     (Error  for  Cephaloon.) 

Typitium  Casey,  Ent.  News,  vol.  9,  p.  193,  1898.  (Genotype,  Cephaloon  ungulate 
LeConte,  Proc.  Boston  Soc.  Nat.  Hist.,  vol.  16,  p.  275,  1873;  original  desig- 
nation and  monobasic.  Placed  in  synonymy  by  Hopping  and  Hopping, 
Pan-Pacific  Ent.,  vol.  10,  p.  64,  1934.) 

Sponidium  Casey,  Ent.  News,  vol.  8,  p.  193,  1898.  (Genotype,  Cephaloon 
tenuicorne  LeConte,  1873;  present  designation.  Placed  in  synonymy  by 
Hopping  and  Hopping,  1934.) 

Ephamillus  Semenow,  Horae  Soc.  Ent.  Ross.,  vol.  34,  p.  495,  1900.  (New 
synonymy.  Genotype,  Cephaloon  variabile  Semenow  «=  Cephaloon  variabilis 
Motschulsky  in  Schrenk,  1860;  original  designation  and  monobasic.) 

Drachylis  Casey,  1898,  Ent.  News,  vol.  9,  p.  195.  (New  synonymy.  Genotype, 
Drachylis  simulans  Casey,  1898;  monobasic.) 

Discussion. — The  type  and  only  known  specimen  of  Drachylis 
simulans  Casey,  the  genotype  of  Drachylis  Casey,  has  been  examined. 
This  is  an  unfortunate  example  of  the  description  of  a  species  on  one 


U.  S.   NATIONAL  MUSEUM 


PROCEEDINGS,   VOL.    103.   PLATE  5 


Adult,  male  genitalia,  and  claws  of  Cephaloidae:  1,  Cephaloon  ienuicorne  LeCorite,  female, 
showing  general  habitus;  2,  paramere  of  male,  C.  tenuicorne;  3,  paramere  of  male,  C. 
bicolor  Horn;  4,  paramere  of  male,  C.  pacificum  Van  Dyke;  5,  paramere  of  male,  C. 
lepturides  Newman;  6,  apical  portion  of  paramere  of  male,  C.  ungulate  LeConte;  7 
abdominal  segments  7-9  of  male,  C.  ungulare;  8,  median  lobe  of  male,  C.  pallens 
Motschulsky;  9,  median  lobe  of  male,  C.  lepturides  Newman;  10,  hind  claw  of  C.  tenui' 
come;  11,  hind  claw  of  C.  pallens;  12,  hind  claw  of  C.  variabilis  Motschulsky. 


BEETLE  FAMILY  CEPHALOIDAE — ARNETT  159 

poor  specimen.  Casey  was  certainly  justified  in  wanting  to  describe 
this  specimen  for  he  beheved  that  it  lacked  the  comblil\:e  claws  charac- 
teristic of  the  other  species  in  the  family,  and  therefore,  its  description, 
even  though  based  on  a  single  specimen,  would  alert  collectors  to  hunt 
for  such  a  strange  cephaloid.  A  close  examination,  however,  reveals  it 
to  have  been  patched,  the  legs  being  undoubtedly  from  some  other 
beetle.  It  appears  to  be  Cephaloon  bicolor  Horn,  with  which  I  synony- 
mize  it. 

The  genus  Ephamillus  Semenow  is  based  ^; on  the  same  variable 
characters  possessed  by  the  other  genera  erected  for  species  of  this 
family.  K6no  (Fauna  Nipponica,  vol.  10,  fasc.  8,  No.  10,  pp.  76-82, 
1937)  illustrates  three  characters,  which,  if  they  were  constant  and  as 
illustrated,  would  serve  for  recognition  of  a  genus.  However,  none  of 
them  appears  to  be  constant  or  as  distinctive  as  thought  by  Kono  and 
others.  The  acute  pulvilli,  curved  at  the  tips,  are  found  in  three 
species,  C.  jpacificum,  C.  ungulare,  and  C.  variabilis.  The  shape  of  the 
pronotum,  as  illustrated  by  Kono  for  C.  variabilis,  is  subject  to  the 
same  sort  of  variation  in  all  the  species.  Finally,  the  sinuation  of  the 
hind  tibia  of  G.  variabilis,  reported  to  be  so  pronounced  that  a  portion 
of  the  tibia  is  thrown  out  of  line  at  least  a  distance  equal  to  the  width 
of  the  tibia,  is  often  barely  perceptible  even  under  a  microscope,  and 
on  some  specimens  of  the  species  it  cannot  be  seen  at  all.  For  these 
reasons,  I  feel  that  this  genus  is  invalid  and  I  place  it  in  synonymy 
with  Cephaloon. 

The  marginate  pronotum  eliminates  the  genus  Stenocephaloon  Pic, 
1932  (Melanges  Exotico-Entomologiques,  fasc.  59,  p.  2;  genotype, 
Stenocephaloon  metallicum  Pic,  monobasic)  from  this  family.  Until 
specimens  can  be  studied,  it  is  best  placed  in  the  family  Serropalpidae 
near  the  genus  Stenotrachelus  Berthold. 

The  following  key  to  the  known  species  of  this  family  is  adapted 
from  Hopping  and  Hopping  (Pan-Pacific  Ent.,  vol.  10,  pp.  64-70, 
1934) .  For  identification  purposes,  the  illustrations  which  accompany 
that  paper  are  very  useful. 

Key  to  the  species  of  Cephaloidae 

1.  Pulvilli  of  tarsal  claws  slender,  acute,  and  curved  at  tips  (pi.  5,  fig.  12) 2 

Pulvilli  of  tarsal  claws  robust,  obtuse,  not  curved  at  tips  (pi.  5,  figs.  10,  11)  _       4 

2.  Three  distal  antenoal  segments  together  approximately   3  mm.   long;  hind 

femora  at  most  simply  curved;  1 0  to  15  mm.  (United  States) 3 

Three  distal  antennal  segments  together  not  over  1.5  mm.  long;  hind  femora 
tending  to  be  sinuate;  size  17  to  20  mm.  (Amur  River,  Eastern  Siberia,  and 
Japan) C.  variabilis  Motschulsky 


160  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  103 

3.  Lateral  margins'of  pronotum  behind  middle_ distinctly  [emarginate;  Western 

United  States C.  pacificum  VanDyke 

Lateral  margins  of  pronotum  behind  middle  almost  straight  (Eastern  United 
States) C.  ungulare  LeConte 

4.  Antennae  with  last  three  segments  thickened 5 

Antennae  with  last  three  segments  not  markedly  thickened 6 

5.  Elytra  without  sutural  and  marginal  black  stripes  (Eastern  United  States) . 

C  lepturides  Newman 

Elytra  with  sutural  and  marginal  black  stripes  (Amur  River,  Eastern  Siberia 

and  Japan) C.  pallens  Motschulsky 

6.  Three  distal  antennal  segments  together  approximately  1.5  mm.  long  (West 

Coast  of  United  States) C.  bicolor  Horn 

Three  distal  antennal  segments  together  barely  over  1  mm.  long  (Western 
United  States  and  Canada) 7 

7.  Male  with  bifurcate  process  of  third  visible  abdominal  sternite  long  and  finger- 

like; female  with  apical  abdominal  sternite  shallowly  and  narrowlj'  emargi- 

nate;  color  of  female  black  to  testaceous C.  tenuicorne  LeConte 

Male  with  bifurcate  process  of  third  visible  abdominal  sternite  shorter,  more 
triangular;  female  with  apical  abdominal  sternite  deeply  emarginate;  color 
of  female  reddish  testaceous  with  elj'tra  and  metasternum  black. 

C.  vandykei  Hopping  and  Hopping 

Key  to  the  species  of  Cephaloon,  based  upon  the  male  genitalia 

Male  of  Cephaloon  variabilis  Motschulsky  unknown. 

1.  Ratio  of  length  of  paramere  to  length  of  basal  piece  not  over  1:2 2 

Ratio  of  length  of  paramere  to  length  of  basal  piece  not  less  than  1:3.5 3 

2.  Median  lobe  tapering  abruptly  at  apical  third,  slender  (pi.  5,  fig.  8) ;  paramere 

lobes  long  and  slender C.  pallens    Motschulsky 

Median  lobe  evenly  tapering  from  base,  more  robust  (pi.  5,  fig.  9) ;  paramere 
lobes  shorter  and  heavier  (pi.  5,  fig.  5) C.  lepturides  Newman 

3.  Paramere  lobes  acute  at  apex  (pi.  5,  fig.  6);  median  lobe  long  and  slender, 

sides  more  parallel C.  ungulare  LeConte 

Paramere  lobes  enlarged  at  apex;  median  lobe  shorter  and  more  robust  at 
apex 4 

4.  Apex  of  paramere  lobes  about  twice  width  of  base  when  viewed  laterally 

(pi.  5,  fig.  3) C  vandykei  Hopping  and  Hopping  and  C.  bicolor  Horn 

Apex  of  paramere  lobes  about  same  width  as  at  base 5 

5.  Apex  of  basal  piece  laterally  extending  beyond  base  of  paramere  to  about 

one-half  length  of  paramere  lobes;  paramere  lobes  stout,  blunt  at  apex, 

uniform  in  width  (pi.  5,  fig.  4) C,  pacificum  VanDyke 

Apex  of  basal  piece  laterally  extending  only  slightly  beyond  base  of  paramere; 
paramere  lobes  more  slender,  and  slightly  enlarged  at  apex  (pi.  5,  fig.  2). 

C.  tenuicorne   LeConte 

In  view  of  the  relatively  recent  and  thorough  discussion  of  the 
species  by  Hopping  and  Hopping  (1934),  I  feel  that  it  is  superfluous 
to  repeat  it  here  except  to  mention  that  under  C.  bicolor  the  name 
Drachylis  simulans  Casey  should  be  added  as  a  synonym  (see  discus- 
sion under  generic  synonymy  preceding) . 


BEETLE    FAMILY    CEPHALOIDAE — AENETT  161 

There  follows  the  bibliographic  citations  of  the  two  exotic  species 
not  mentioned  by  Hopping  and  Hopping.  The  key  and  illustrations, 
I  believe,  sufficiently  characterize  them.  A  revised  section  of  the 
Coleopterorum  Catalogus  (Junk)  is  being  prepared  which  will  give  all 
of  the  literature  references,  therefore  they  will  not  be  cited  here. 

Cephaloon  pallens  Motschulsky,  1860 

Plate  5,  Figures  8,  11 

Cephaloon  pallens  Motschulsky,  in  Schrenck,  Reisen  und  Forschungen  in  Amur- 
lande,  vol.  2,  pt.  2,  p.  140,  pi.  9,  fig.  15,  1860.  (Type  locality,  Kisi,  on  the 
River  Amur.) 

Cephaloon  pallens  var.  cindipennie  Heyden,  Deutsche  Ent.  Zeitschr.,  p.  167. 
1892.  (Type  locality  unknown,  except  "Amur"  as  given  in  the  title  of  the 
paper.) 

Cephaloon  pallens  var.  koltzei  Heyden,  Deutsche  Ent.  Zeitschr.,  p.  168,  1892. 
(Type  locality,  "Amur,"  as  above.) 

Cephaloon  pallens  var.  maculicolle  Heyden,  Deutsche  Ent.  Zeitschr.,  p.  167,  1892. 
(Type  locality,  "Amur,"  as  above.) 

Cephaloon  pallens  var.  picticolle  Heyden,  Deutsche  Ent.  Zeitschr.,  p.  167,  1892. 
(Type  locality,  "Amur,"  as  above.) 

1  Cephaloon  pallens  Motschulsky,  Solsky,  Horae  Soc.  Ent.  Rossicae,  vol.  11,  p. 
295,  1875.  (Djalinda  River.  Note:  This  is  probably  a  misidentification  of 
Ephamillus  variabilis  var.  tristiculus  Heyden,  1892,  judging  from  the  descrip- 
tion and  locality.) 

Cephaloon  sakurae  Lewis,  Ann.  Mag.  Nat.  Hist.,  ser.  6,  vol.  15,  p.  444,  fig.  10, 
1895.  (Placed  as  synonym  by  K6no,  1937,  Fauna  Nipponica,  vol.  10,  fasc. 
8,  No.  10,  p.  79.) 

In  a  species  with  the  extreme  color  variation  exhibited  in  all  of  the 
species  of  this  family,  the  naming  of  a  few  of  the  color  variants  serves 
no  useful  purpose  and  these  are  therefore  here  disregarded. 

Cephaloon  variabilis  Motschulsky,  1860 

Plate  5,  Figure  12 

Cephaloon  variabilis  Motschulsky,  in  Schrenck,  Reisen  und  Forschungen  in  Amur- 
lande,  vol.  2,  pt.  2,  p.  141,  1860.  (Type  locality,  Marunsk  on  the  River 
Amur.) 

Cephaloon  variabilis  var.  tristiculus  Heyden,  Deutsche  Ent.  Zeitschr.,  1892,  p.  169. 
(Type  locality,  unknown  except  "Amur.") 

Here  again,  the  naming  of  a  single  color  form  can  serve  no  useful 
purpose. 


II,  S.  GOVERNMENT  PRINTING  OFFICE:  1953 


PROCEEDINGS  OF  THE   UNITED  STATES   NATIONAL  MUSEUM 


SMITHSONIAN  INSTITUTION 

U.  S.  NATIONAL  MUSEUM 


Vol.   103  Washington  :  1953  No.  3322 


THE  FRESH-WATER  TRICLADS  (TURBELLARIA)  OF  ALASKA 


By  Roman  Kenk 


Introduction 


In  1948,  during  an  investigation  of  biting  insects  in  Alaska  con- 
ducted by  the  Bureau  of  Entomology  and  Plant  Quarantine,  United 
States  Department  of  Agriculture,  Dr.  Reese  I.  SaUer  collected  and 
transmitted  to  me  several  samples  of  fresh-water  triclads  (planarians) . 
An  examination  of  the  material  revealed  that  the  worms  belonged  to 
the  genus  Polycelis,  a  genus  fairly  common  in  Asia  and  Europe  but 
only  twice  reported  from  North  America.  The  finding  suggested  a 
closer  relationship  of  the  Alaskan  fresh-water  fauna  with  that  of  East 
Asia.  A  more  thorough  study  of  the  triclads  of  Alaska  promised  to 
yield  a  more  definite  understanding  of  these  zoogeographical  relations. 

My  field  trip  ^  to  Alaska  was  made  in  the  sunmaer  of  1950.  Since 
the  available  time  was  rather  limited,  the  collecting  was  done  mainly 
along  the  highways  of  the  territory  and  in  the  vicinities  of  Point 
Barrow  and  Umiat. 

The  following  roads  and  fresh-water  localities  were  visited  on  the 
trip:  Steese  Highway  in  the  section  between  Fairbanks  and  Birch 
Creek  (milepost  103);  Elliot  Highway,  from  Fairbanks  to  Livengood; 
tundra  lakes  and  pools  in  the  vicinity  of  Point  Barrow;  ColvUle 
River,  several  streams,  and  a  lake  near  Umiat;  Mount  McKinley 
Park  Road,  a  section  of  about  15  miles  adjoining  the  railroad  station; 
Glenn  Highway,  from  Anchorage  to  Glenallen;  road  from  Palmer  to 

'  This  study  was  supported  by  the  Arctic  Institute  of  North  America  under  contractual  arrangement 
with  the  OfBce  of  Naval  Research. 

238538—53 1  163 


164  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  los 

Willow;  road  from  Anchorage  to  Potter;  and  Richardson  Highway, 
section  between  Valdez  and  Glenallen. 

No  fresh-water  triclads  were  found  by  me  either  at  Point  Barrow 
or  at  Umiat.  I  received,  however,  from  the  U.  S.  National  Museum, 
a  sample  of  triclads  collected  by  P.  F.  Scholander  in  a  lake  near  Umiat. 
The  relative  scarcity  of  planarians  in  tundra  lakes  and  pools  may  be 
due  to  the  high  acidity  of  the  waters  or  to  the  extremely  severe  con- 
ditions prevailing  in  them  during  the  long  winter  season. 

Grateful  acknowledgment  is  made  of  the  very  helpful  cooperation 
which  was  extended  to  me  by  several  agencies  in  Alaska:  The  Arctic 
Research  Laboratory,  Point  Barrow;  the  U.  S.  Public  Health  Service, 
Anchorage;  the  Alaska  Road  Commission,  Fairbanks,  Anchorage,  and 
Glenallen;  and  the  Division  of  Forestry,  Bureau  of  Land  Management, 
Fairbanks  and  Anchorage.  I  also  wish  to  express  my  indebtedness  to 
Prof.  Edward  G.  Reinhard  of  the  Catholic  University  of  America  and 
to  Dr.  Fenner  A.  Chace,  Jr.,  and  Dr.  Doris  M.  Cochran  of  the  U.  S. 
National  Museum  who  kindly  permitted  me  to  use  their  laboratory 
and  office  facilities  in  Washington,  D.  C. 

Four  species  of  fresh-water  triclads  were  collected  in  Alaska.  Two 
are  inhabitants  of  the  White  Mountains,  a  mountain  range  extending 
in  an  east-west  direction  north  of  Fairbanks.  The  other  two  are 
widely  distributed  in  waters  of  the  Alaska  Range  and  of  the  southern 
section  of  Alaska  and  one  of  them  reaches  as  far  north  as  Umiat. 

Family  Planariidae 
Genus  Phagocata  Leidy 

Phagocata  niveau  new  species 

Figure  21;  Plate  6,  Figure  1 

Description. — This  is  a  slender,  rather  delicate  species.  Mature 
specimens  measure  up  to  8  mm.  in  length  and  about  L5  mm.  in  width. 
In  the  quietly  gliding  animal  the  anterior  end  is  truncated,with  a  very 
slightly  bulging  frontal  outline  and  with  rounded  lateral  corners 
(auricles) .  There  is  no  distinct  narrowing  or  neck  behind  the  auricles 
and  the  lateral  margins  of  the  head  are  approximately  parallel.  Behind 
the  head,  the  body  widens  and  soon  reaches  its  greatest  width.  From 
there  on,  the  lateral  margins  of  the  body  run  parallel  up  to  the  level 
of  the  mouth,  to  converge  agam  in  the  postpharyngeal  region  and  to 
meet  in  a  bluntly  pointed  posterior  end. 

The  species  lacks  pigment  and  usually  appears  white  and  somewhat 
transparent.  The  intestinal  contents  may  shine  through  the  body 
wall  and  give  the  animal  a  certain  amount  of  color;  the  margins  of  the 
body,  the  head  region,  and  the  areas  occupied  by  the  pharynx  and  the 
copulatory  apparatus,  however,  are  always  white. 


FRESH-WATER   TRICLADS    OF   ALASKA — ^KENK  165 

There  are  two  rather  small  eyes,  situated  close  together  (about  one- 
fourth  the  body  width  apart  at  the  level  of  the  eyes)  and  far  removed 
from  the  frontal  end.  This  character,  easily  recognized  in  life,  dis- 
tinguishes the  species  from  another  white  triclad  with  which  it  shares 
its  habitat,  Dendrocoelopsis  alaskensis,  described  as  new  on  p.  178. 

The  pharynx  is  inserted,  in  sexually  mature  specimens,  somewhat 
behind  the  middle  of  the  body  and  measures  about  one-sixth  of  the 
body  length.  The  copulatory  organs  occupy  the  anterior  half  of  the 
postpharyngeal  region. 

The  animal  moves  by  gliding  only;  crawling  movements,  such  as 
are  seen  in  other  triclads,  particularly  in  those  equipped  with  anterior 
adhesive  organs,  have  not  been  observed  in  this  species. 

From  the  description  it  may  be  seen  that  the  species  in  life  shows 
a  close  resemblance  to  other  species  of  the  same  genus,  particularly 
to  the  American  Phagocata  morgani,  the  European  P.  albissima,  P. 
vitta,  and  related  forms.  A  separation  of  these  species  can  be  made 
only  on  the  basis  of  anatomical  characters. 

Only  those  characters  of  the  digestive  system  that  have  a  taxonomic 
significance  are  discussed  here.  The  pharynx  has  a  structure  typical 
of  the  family  Planariidae;  i.  e.,  the  fibers  of  the  internal  muscle  zone 
are  arranged  in  two  distinct  layers,  a  thick  inner  circular  layer  and  a 
narrower  outer  longitudinal  one.  The  anterior  intestinal  trunk  bears 
10  or  11  branches  on  each  side.  Each  posterior  trunk  has  21  to  27 
lateral  branches  and  numerous  short  medial  branches  in  both  the 
pharyngeal  and  postpharyngeal  regions. 

The  testes  are  numerous  and  are  arranged,  on  each  side  of  the 
midline,  in  a  longitudinal  zone  extending  from  a  short  distance  behind 
the  ovary  almost  to  the  posterior  end  of  the  body.  Their  position  is 
predominantly  ventral,  below  the  intestinal  branches.  Only  a  few 
testes  extend  into  the  mesenchymatic  "septa"  between  the  branches 
toward  the  dorsal  side. 

The  two  ovaries  are  typical,  each  situated  approximately  below  the 
second  intestinal  branch.  An  undifferentiated  mass  of  cells,  the  par- 
ovarium, is  attached  to  the  dorsolateral  side  of  each  ovary. 

The  genital  pore  (pg),  situated  about  halfway  between  the  mouth 
and  the  posterior  end  of  the  body,  leads  into  a  small  cavity  (ac) 
which  continues  to  the  left  and  dorsally  into  the  duct  of  the  copulatory 
bursa  (bd)  and  to  the  right  and  anteriorly  into  the  male  atrium  (am) . 
This  cavity  may  be  considered  to  represent  a  common  genital  atrium. 
In  some  specimens,  however,  there  appears  to  be  no  differentiation 
of  the  atrium  into  male  and  common  parts,  and  the  bursa  duct  and 
an  undivided  atrium  meet  at  or  near  the  genital  pore.  These  varia- 
tions are  obviously  due  to  the  different  states  of  muscular  contraction 
in  which  the  animals  were  killed.  The  atrium,  narrow  at  the  genital 
aperture,  widens  as  it  extends  forward,  to  the  right  side  of  the  midline. 


166  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  io3 

It  is  lined  with  a  tall,  glandular  epithelium,  the  cells  of  which  project 
into  the  cavity  in  a  villuslike  fashion.  Below  the  epithelium  there 
are  two  layers  of  muscle  fibers,  one  circular  and  the  other  longitudinal. 

The  penis  consists  of  a  spherical,  muscular  bulb  embedded  in  the 
mesenchyme,  and  a  moderately  large  papilla  projecting  into  the  male 
atrium.  The  bulb  is  pierced  by  numerous  gland  ducts  which  open 
into  the  lumen  of  both  the  bulb  and  the  papilla.  The  shape  of  the 
papilla  is  subject  to  great  variation,  due  apparently  to  the  state  of 
contraction  of  the  organ.  It  may  be  twisted  to  one  side  and  even 
partly  inverted  into  the  lumen  of  the  penis  (similar  to  the  pseudo- 
flagellum  of  various  dendrocoelids) .  The  shape  shown  in  figure  21 
appears  to  be  that  of  the  organ  at  rest.  The  outer  wall  of  the  papilla 
is  covered  with  a  tall  to  cubical  epithelium  similar  to  that  lining  the 
atrium.  Below  the  epithelium  there  is  a  layer  of  circular  muscle 
fibers  followed  by  another  of  longitudinal  fibers.  The  shape  of  the 
penis  lumen  {Ip)  is  as  changeable  as  that  of  the  papilla.  Typically, 
it  appears  to  be  wider  in  the  bulb  than  it  is  in  the  papilla,  though 
there  is  no  distinct  ejaculatory  duct  differentiated.  The  lumen  opens 
ventrally  to  the  tip  of  the  papUla.  The  two  vasa  deferentia  {vd) 
penetrate  the  penis  bulb  from  both  sides  and  empty  into  the  penis 
lumen  separately,  but  not  far  apart. 

The  two  oviducts  converge  at  the  level  of  the  copulatory  apparatus, 
the  left  one  passing  between  the  bursa  duct  and  the  male  atrium,  and 
unite  at  a  point  dorsally  to  the  atrium.  The  rather  long  common 
oviduct  {ode)  curves  ventrally  and  opens  into  the  posterior  part  of  the 
male  atrium.  The  terminal  sections  of  the  paired  oviducts  and  the 
greater  part  of  the  common  oviduct  receive  the  outlets  of  numerous 
eosinophilic  glands,  the  cell  bodies  of  which  are  scattered  in  the  sur- 
rounding mesenchyme,  particularly  dorsally  to  the  atrium. 

The  copulatory  bursa  (6)  is  of  moderate  to  large  size  and  is  ir- 
regularly lobed.  The  bursa  duct  or  stalk  {hd)  is  wide,  runs  posteriorly 
to  the  left  of  the  midline,  and  curves  ventrally  to  reach  the  genital 
aperture.  It  is  lined  with  a  tall,  glandular  epithelium  and  surrounded 
with  a  strong  muscular  coat  consisting  of  intermingled  circular  and 
longitudinal  fibers. 

Taxonomic  position. — The  genus  Phagocata  Leidy  (Fonticola 
Komarek)  in  its  present  extent  (cf.  Hyman,  1937,  pp.  300-302)  has 
representatives  in  Europe,  Asia,  and  North  America.  The  genus  is 
not  quite  homogeneous  and  will  probably,  in  due  time,  be  subdivided 
into  several  genera  (cf.  Beauchamp,  1939).  For  the  purpose  of  com- 
parison, we  may  consider  here  only  those  species  of  the  genus  that 
lack  pigment.  Though  the  presence,  or  the  lack,  of  pigment  is  a 
character  of  subordinate  taxonomic  value,  it  may  nevertheless  serve 
well  as  a  character  of  specific  rank.     The  Alaskan  form  differs  from 


FRESH-WATER    TRICLADS    OF    ALASKA — ^KENK 


167 


European  species  such  as  P.  vitta  (Duges)  and  P.  albissima  (Vejdovsk^) 
and  from  the  American  white  P.  morgani  (Stevens  and  Boring)  mainly 
in  the  structm-e  of  the  male  copulatory  organ.  Asia  has  several 
species  of  Phagocata,  the  majority  of  them  pigmented  forms.  Of  the 
three  unpigmented  Asiatic  species  that  may  belong  to  the  genus,  two 


Figure  21. — Phagocata  nivea,  diagram  ot  the  copulatory  organs  In  longitudinal  section, 
X  92.  ac,  common  atrium;  am,  male  atrium;  b,  copulatory  bursa;  bd,  bursa 
stalk;  Ip,  penis  lumen;  o,  mouth;  ode,  common  oviduct;  pg,  genital  pore;  vd,  vas  deferens. 


have  been  described  from  immature  specimens  and  the  anatomy  of 
their  reproductive  systems  is  not  known:  Planaria  pellucida  Ijima 
and  Kaburaki  (1916)  from  Sakhalin  and  a  species  from  the  Bailial 
region  assigned  tentatively  to  Fonticola  by  Bazikalova  (1947).  A 
third  species,  Phagocata  coarctata  (Arndt,  1922),  from  the  vicinity  of 
Vladivostok,  is  sufficiently  well  Imown,  although  no  fully  mature 
individuals  of  this  species  have  been  studied.  P.  coarctata  differs 
from  P.  nivea  externally  in  being  smaller  and  broader,  and  in  having 
a  different  contour  of  the  anterior  end,  which  bears  protruding  lateral 
lobes,  and  a  greater  distance  between  the  two  eyes.  Anatomically, 
the  two  species  are,  undoubtedly,  closely  related.^ 


'  Livanov  and  Zabusova  (1940,  p.  146)  state  that  a  reexamination  of  Amdt's  slides  of  PUmaria  coarctata 
showed  that  the  arrangement  of  the  muscle  fibers  of  the  pharynx  conformed  with  the  dendrocoelid  type 
(circular  and  longitudinal  fibers  of  the  internal  muscle  zone  intermingled).  Amdt  (1922,  p.  108)  described 
the  anatomy  of  the  pharynx  in  minute  detail  and  indicated,  both  in  a  figure  (pi.  4,  fig.  7)  and  in  the  text, 
a  typical  planariid  pattern.  I  must  assume  that  some  confusion  occurred  somewhere,  probably  in  the 
identification  of  the  slides  sent  to  Livanov. 


168  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  io3 

In  Phagocata  nivea  the  penis  lumen  opens  usually  below  the  tip  of 
the  penis  papilla.  This  character  has  been  used  by  Livanov  and 
Zabusova  (1940,  p.  96)  to  segregate  a  group  of  Asiatic  species  from 
Phagocata  and  to  place  them  in  a  new  genus,  Penecurva.  The  same 
character  is  found,  however,  in  a  common  North  American  species, 
Phagocata  morgani,  which  shows  no  other  close  relations  with  the 
Asiatic  group.  This  character  is  apparently  inadequate  as  a  basis 
for  the  establishment  of  a  new  genus. 

Holotype. — On  one  slide,  USNM  22332,  creek  crossing  EUiot  High- 
way at  milepost  ^31.0,  July  24,  1950. 

Distribution  and  ecology. — Phagocata  nivea  was  collected  in  cool, 
fast  mountain  streams  on  the  slopes  of  the  White  Mountains,  a  range 
north  of  Fairbanks.  The  temperature  of  the  water  ranged  from  3.2 
to  7.2°  C.  (July).  The  animals  are  cold-stenothermic  and  do  not 
tolerate  sudden  increases  in  temperature.  They  were  often  found 
in  the  company  of  another  white  triclad,  Dendrocoelopsis  alaskensis. 

Stream  on  Steese  Highway  (pi.  8),  at  milepost  84.0,  altitude  2,700  feet,  July 
19  and  21,  1950,  water  temperature  6.9°  C;  one  immature  and  one  mature  speci- 
men, from  under  stones. 

Willow  Creek,  on  Steese  Highway,  at  milepost  96.6,  altitude  2,100  feet,  July 
19,  1950,  7.2°  C;  two  immature  specimens,  from  under  stones. 

Spring  and  stream  on  Steese  Highway,  milepost  82.5,  near  Alaska  Road  Com- 
mission camp,  July  21,  1950;  two  mature  and  six  immature  specimens,  on  the 
undersides  of  stones. 

Fox  Gulch,  on  Elliot  Highway,  milepost  1.2,  July  24,  1950,  6.0°  C,  one  small 
specimen  near  bait  (beef  liver). 

Creek  crossing  Elliot  Highway  at  milepost  31.0,  July  24,  1950;  39  specimens, 
about  half  of  them  mature,  collected  under  stones  (holotype). 

Genus  Polycelis  Ehrenberg 
Polycelis  borealis^  new  species 

Figure  22;  Plate  6,  Figure  2 

Description. — Mature  animals  are  usually  12  to  15  mm.  long  and 
1.5  to  2  mm.  wide  (larger  specimens,  measuring  up  to  20  mm.  in 
length,  have  been  seen).  The  frontal  margin  is  slightly  convex.  At 
the  lateral  corners  of  the  head  there  is  a  pair  of  elongated,  bluntly 
pointed  auricles  which  are  held  raised  when  the  animal  is  gliding 
quietly.  Behind  the  auricles,  the  body  first  narrows  slightly,  then 
gradually  widens,  reaching  its  greatest  width  at  the  level  of  the 
pharynx.  Behind  the  pharynx  the  body  tapers  to  a  moderately 
pointed  posterior  end. 

The  color  of  the  dorsal  side  is  usually  a  uniform  light  or  dark 
brown,  that  of  the  ventral  side  a  light  grayish  brown.     In  animals 

'  The  large  highways  of  Alaska  are  marked  with  wooden  mileposts  indlcatmg  only  full  miles.  Many 
posts  were  missing  at  the  time  when  the  collections  were  made.  Fractional  milages  were  usually  estimated 
from  the  nearest  milepost  or  from  the  speedometer  readings  of  the  car  used. 


FRESH-WATER    TRICLADS    OF   ALASKA — ^KENK  169 

from  some  localities,  an  indistinct  lighter  median  line  occurred  dor- 
sally  in  the  prepharyngeal  region,  with  lighter  areas  above  the  pharynx 
and  the  copulatory  organs. 

The  species  has  many  small  eyes  (a  generic  character)  arranged 
in  a  band  along  the  frontal  margin  of  the  head,  the  base  of  the  auricles, 
and  the  lateral  margins  of  the  body  a  short  distance  behind  the  head. 
Anteriorly  the  eyes  are  placed  in  more  than  one  row,  somewhat 
irregularly  scattered ;  behind  the  head  they  are  in  a  single  row  reaching 
backward  about  one-third  to  one-half  the  length  of  the  prepharyngeal 
region.  There  is  no  narrowing  of  the  band  of  eyes  at  the  base  of  the 
auricles. 

The  pharynx  is  inserted  at  about  the  middle  of  the  body  and 
measures  in  length  almost  one-fourth  the  length  of  the  body.  The 
copulatory  organs  occupy  more  than  half  the  post-pharyngeal  region. 

The  animal  moves  by  gliding  only. 

The  pharynx  of  Polycelis  borealis  is  structurally  typical  of  the  genus 
Polycelis  and  of  the  family  Planariidae,  the  muscle  zone  being  formed 
by  two  distinct  layers,  a  thick  circular  layer  adjoining  the  epithelium 
of  the  pharyngeal  lumen  and  a  thinner  layer  of  longitudinal  fibers. 
The  anterior  trunk  of  the  intestine  bears  5  to  6  lateral  branches. 

The  numerous,  fairly  large  testes  are  arranged  in  two  zones,  to 
the  right  and  left  of  the  anterior  intestinal  trunk,  extending  from  the 
level  of  the  ovaries  posteriorly  to  the  base  of  the  pharynx.  The 
testes  are  essentially  ventral,  though  individual  vesticles  may  extend 
dorsally  in  the  mesenchymatic  spaces  between  the  intestinal  branches 
and  occupy  almost  the  entire  dorsoventral  diameter  of  the  body. 
Where  intestinal  branches  are  present,  however,  the  testes  develop 
only  below  them.  Each  zone  of  testes  reaches  laterally  only  little 
beyond  the  ventral  nerve  cord. 

The  ovaries  are  situated  behind  the  first  pair  of  lateral  branches  of 
the  anterior  trunk  of  the  intestine. 

The  genital  pore  (j)g),  situated  in  the  midline  behind  the  middle  of 
the  postpharyngeal  region,  connects  with  a  narrow  posterior  extension 
of  the  genital  atrium  (a) .  This  extension  leads  dorsally  and  somewhat 
to  the  left  into  the  duct  of  the  copulatory  bursa  and  anteriorly  into 
the  widened  portion  of  the  atrium  containing  the  penis.  There  is  no 
marked  division  of  the  atrium  into  a  posterior  common  atrium  and 
an  anterior  male  atrium.  The  wall  of  the  atrium  is  lined  with  a 
cubical  epithelium  and  equipped  with  two  muscle  layers,  one  circular 
and  the  other  longitudinal. 

The  penis  consists  of  a  bulb  embedded  in  the  mesenchyme  and  a 
papUla  projecting  into  the  atrium.  Neither  of  the  two  parts  is  very 
muscular.     The  shape  of  the  penis,  particularly  of  the  papilla,  is  very 


170  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  i03 

changeable.  In  specimens  that  were  fixed  in  a  well-extended  condi- 
tion, the  bulb  appears  spherical  and  the  papilla  has  a  conical  shape. 
The  bulb  contains  a  spacious,  irregularly  lobed  cavity,  the  seminal 
vesicle  (vs).  Numerous  gland  ducts  empty  into  the  lumen  of  the 
bulb  after  entering  the  bulb,  particularly  from  its  anterior  surface, 
and  penetrating  between  its  muscle  fibers.  The  ducts  are  filled 
with  a  granular,  slightly  eosinophilic  secretion.  The  two  vasa 
deferentia  (vd)  open  into  the  seminal  vesicle  separately  from  the 
anterolateral  sides. 

The  penis  papUla  is  conical  or  finger-shaped  when  well  extended.  It 
is  covered  with  a  cubical  epithelium  below  which  there  are  two  rather 
feebly  developed  muscle  layers,  tapering  in  thickness  toward  the  tip 
of  the  papilla:  a  layer  of  circular  fibers  adjoining  the  epithelium  and 
a  thinner  layer  of  longitudinal  ifibers.  The  absence  of  a  strong  mus- 
cular wall  probably  accounts  for  the  great  variation  in  the  shape  of 
the  penis  papilla  observed  in  the  material.  The  lumen  of  the  seminal 
vesicle  continues  into  the  papilla  as  a  wide  canal  (de),  opening  at  or 
near  (ventrally  to)  the  tip  of  the  papUla.  The  canal  is  lined  with  an 
epithelium  of  cubical  cells  and  lacks  the  gland  openings  character- 
istic of  the  seminal  vesicle.  It  corresponds  to  an  ejaculatory  duct 
but  it  apparently  has  no  distinct  muscle  coat. 

The  two  oviducts,  riuming  posteriorly  along  the  ventral  nerve 
cords,  bend  dorsally  and  medially  at  the  level  of  the  copulatory 
organs.  The  left  one  passes  through  the  space  between  the  atrium 
and  the  bursa  duct  and  unites  with  the  oviduct  of  the  right  side 
dorsally  to  the  atrium.  The  common  oviduct  {ode)  runs  ventrally 
and  opens  into  the  posterior  part  of  the  atrial  cavity.  The  terminal 
sections  of  the  paired  oviducts  and  a  section  of  the  common  oviduct 
are  equipped  with  strongly  eosinophilic  shell  glands. 

The  copulatory  bursa  (6)  is  a  large  sac  with  a  lobed  outline.  It 
connects  posteriorly  with  a  duct  of  almost  uniformly  wide  diameter, 
the  bursa  stalk  (bd).  The  duct  runs  first  posteriorly,  to  the  left  of 
the  penis  bulb,  then  curves  ventrally  and  opens,  from  the  dorsal  side, 
into  the  narrow  terminal  portion  of  the  atrium,  close  to  the  genital 
pore.  The  bursa  stalk  has  a  strong  coat  of  intermingled  circular  and 
longitudinal  muscle  fibers.  There  is  no  histological  differentiation 
into  anterior  and  posterior  sections  of  the  bursa  stalk. 

Taxonomic  position. — Polycelis  borealis  is  the  second  species  of  the 
genus  Polycelis  to  be  found  on  the  North  American  continent.  The 
other  species,  P.  coronata  (Girard),  reported  from  Wyoming  and  South 
Dakota  (Hyman,  1931),  resembles  it  closely  in  external  appearance 
and  probably  cannot  be  distinguished  from  it  in  life.  In  both  species, 
the  shape  of  the  anterior  end  is  very  similar.  The  auricles  of  P. 
borealis  are  perhaps  a  trifle  longer  and  more  pointed  than  those  of  P. 


U.   S.    NATIONAL   MUSEUM 


PROCEEDINGS,   VOL.    103,    PLATE  6 


Sketches  of  living  specimens  of  the  four  Alaskan  triclad  species,  all  XIO:  1.  Phagocata 
nivea;  2.  Polycelis  borealis;  3.  Dendrocoelopsis  piriformis,  striped  form;  4.  Dendrocoelop- 
sis  alaskensis. 


FRESH-WATER    TRICLADS    OF    ALASKA — KENK 


171 


coronata.  Hyman  (1931,  p.  126)  states  that  in  P.  coronata  the 
curved  band  of  eyes  narrows  as  it  crosses  the  base  of  the  auricles; 
no  such  narrowing  has  been  seen  in  P.  borealis.  These  differences 
are,  however,  insignificant  and  do  not  permit  a  clear  separation  of 
the  two  forms.  The  same  may  be  said  of  other  species  of  the  genus, 
which  likewise  could  be  confused  with  the  American  forms  in  life: 
the  Japanese  species,  P.  auriculata  and  P.  karafto;  and  P.  schmidti 
occurring  both  in  Kamchatka  and  in  Japan.  These  forms  differ 
from  each  other  more  clearly  in  their  anatomical  characters. 

bd     ode 


Figure  22. — Polycelis  borealis,  diagram  of  the  copulatory  organs  in  longitudinal  section, 
X  70.  a,  atrium;  b,  bursa;  bd,  bursa  stalk;  de,  ejaculatory  duct;  o,  mouth;  ode,  common 
oviduct;  pg,  genital  pore;  vd,  vas  deferens;  vs,  seminal  vesicle. 

To  gain  a  better  insight  into  the  systematic  relations  of  Poly- 
celis borealis  to  other  species  of  the  same  genus,  it  is  necessary  to  review 
the  present  state  of  the  systematics  of  the  genus.  The  genus  Poly- 
celis has  a  muscular  pattern  of  the  pharynx  typical  of  the  family 
Planariidae;  the  oviducts  unite,  without  embracing  the  bursa  stalk, 
to  form  an  unpaired  terminal  oviduct;  the  testes  are  situated  in  the 
anterior  part  of  the  body  only;  the  male  atrium  is  not  surrounded 
by  radial  muscle  plates;  and  the  eyes  are  numerous.  The  genus 
thus  defined  (Kenk,  1930)  has  today  about  twenty  species.  It  may 
be  subdivided  into  subgenera  on  the  basis  of  structural  characters 
of  the  copulatory  apparatus. 

Subgenus  Polycelis,  lacking  adenodactyls  and  lacking  an  excessive 
development  of  the  muscle  coat  of  the  male  atrium.  This  subgenus 
includes  Sorocelides  Sabussowa  (1929,  p.  521)  and  Polycelidia  Zabu- 
sova  (1936,  p.  152),  both  described  as  distinct  genera.  The  subgenus 
comprises  the  following  species: 


238538—53 


172  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  103 

Polycelis  nigra  (O.F.MulleT),  Europe  P.    polyopis    Zabusova    (1936),    Kam- 

P.  receptaculosa  (Livanov  and  Zabusova,  chatka 

1940),  Teletskoe  Lake  in  the  Altai  P.  karafto  Ijima  and  Kaburaki  (1916), 
Mountains  Sakhalin 

P.    eburnea    (Muth,     1912),    Aral    Sea  P.  sopporo  (Ijima  and  Kaburaki,  1916), 
region  Japan 

P.  ttfeeftca  Hyman  (1934),  Tibet  P.   coronata    (Girard,    1894),    Wyoming 

P.  koslowi  (Zabusov,  1911),  Tibet  and  South  Dakota 

P.   elongata    (Sabussowa,    1929),   Kam- 
chatka 

Subgenus  Seidlia  Zabusov  (1911,  suppl.,  p.  7),  distinguished  by  an 
extraordinarily  thick  muscle  zone  surrounding  the  male  atrium. 
Zabusov's  (1916,  p.  273)  genus  Rjabuschinskya  likewise  belongs  here. 
Species  of  the  subgenus: 

Polycelis    sabussowi    (Seidl,    1911),    in-  P.  schmidti  (Zabusov,  1916),  including 
eluding     Seidl's     species     Sorocelis  P.  ijimai  Kaburaki  (1922),  Kam- 

sabussowi,   S.  lactea,    S.    stummeri  chatka,  Kurile  Islands,  and  Japan 

(cf.   Kenk,  1936),  Turkestan  P.  auncu/a<a  Ijima  and  Kaburaki  (1916), 

P.  relicta  (Sabussowa,  1929),  Kamchatka  Japan 

P.   eurantron   (Zabusova,   1936),   Kam- 
chatka 

The  third  subgenus,  Ijimia  Bergendal  (1890,  p.  326),  is  charac- 
terized by  possessing  solid  adenodactyls : 

Polycelis    felina     (Dalyell),     including  P.  linkoi  Zabusov  (1901),  Onega  Lake 
P.    cornuta  Johnson,    Europe   and  in  European  Russia 

North  Africa  P.  oculi-marginata  (Palombi,  1931),  New 

P.   tenuis  Ijima,   Europe  and  western  Guinea  (see  Beauchamp,  1947) 

Asia 

Two  species  of  the  genus  Polycelis  are  too  mcompletely  known  to 
permit  their  assignment  to  either  the  subgenus  Polycelis  or  Seidlia: 
P.  eudendrocoeloides  (Sabussowa,  1929)  from  the  Kamchatka  Penin- 
sula and  P.  tibetica  (Zabusov,  1911,  p.  349)  from  Tibet  (P.  tibetica 
Hyman,  1934,  will  have  to  be  renamed  if  it  should  prove  to  be  different 
from  Zabusov's  species) . 

Polycelis  borealis  is  clearly  a  member  of  the  subgenus  Polycelis. 
It  differs  from  P.  nigra  in  having  weU-developed  auricles;  from  P. 
elongata,  P.  polyopis,  and  P.  sapporo,  in  the  arrangement  of  the  eyes; 
and  from  P.  coronata  and  P.  receptaculosa  in  the  structure  of  the  male 
copulatory  organ.  The  reproductive  system  of  P.  karafto  has  not  been 
described  adequately  and  thus  does  not  permit  a  comparison  with 
that  of  the  Alaskan  species.  The  remaining  species  of  the  subgenus, 
P.  eburnea,  P.  koslowi,  and  P.  tibetica  Hyman,  two  of  which  are  known 
only  from  the  study  of  preserved  specimens,  agree  with  P.  borealis 
in  being  pigmented  forms  with  prominent  auricles  and  in  having 
a  similar  anatomy  of  the  copulatory  organs.     The  four  species  are 


FRESH-WATER    TRICLADS    OF   ALASKA — KENK  173 

undoubtedly  very  closely  related;  whether  some  of  them  are  identical 
or  are  the  same  as  the  Alaskan  form  cannot  be  established  on  the  basis 
of  present  knowledge.  The  Alaskan  Poly  cells,  therefore,  is  described 
as  a  new  species, 

Holotype. — On  three  slides,  USNM  22333,  clear  spring  on  the  road 
from  Palmer  to  Willow,  20.1  mUes  from  Palmer,  altitude  3,800  feet, 
Aug.  9,  1950. 

Distribution  and  ecology. — Polycelis  borealis  is  a  very  common  species 
occurring  in  mountain  streams  in  the  southern  part  of  Alaska  (Alaska 
Range,  Talkeetna  Mountains,  Chugach  Range).  Typically  it 
inhabits  clear,  cold,  fast-running  waters.  It  has  also  been  found, 
however,  in  several  small  mountain  lakes  which  connect  with  streams 
in  which  the  species  lives.  It  has  not  been  observed  in  sUt-bearing 
glacier  streams. 

The  typical  temperature  range  of  the  habitats  was  between  3.0° 
and  about  15°  C.  (August  1950).  Temperatures  above  15°  C.  (up 
to  22.4°  C.)  were  encountered  only  rarely  and  only  in  habitats  which 
presumably  have  great  diurnal  temperature  amplitudes  (shallow  lakes, 
small  exposed  streams). 

The  great  majority  of  the  specimens  collected  was  asexual.  Sexually 
mature  animals  were  seen  in  only  6  of  the  31  localities  in  which  the 
species  was  found.  In  some  localities,  a  large  percentage  of  the  speci- 
mens exhibited  freshly  regenerated  heads  or  posterior  ends,  indicating 
that  vivid  asexual  reproduction  was  taking  place. 

Clear  springs  on  the  road  from  Palmer  to  Willow,  20.1  miles  from  Palmer, 
altitude  3,800  feet,  Aug.  9,  1950,  water  temperature  3.0°  C,  numerous  specimens, 
five  mature  (holotype). 

Clear,  fast  mountain  stream  crossing  Mount  McKinley  Park  road,  5.5  miles 
from  the  railroad  station;  Aug.  5,  1950. 

Streams  on  road  from  Palmer  to  Willow  11.9,  18.0,  19.2,  20.2,  21.6,  22.1,  22.8, 
and  34.0  miles  from  Palmer,  and  Ice  Lake,  20.4  miles  from  Palmer,  Aug.  9,  1950. 

Streams  on  Glenn  Highway  (Anchorage  to  junction  with  Richardson  Highway 
near  Glenallen)  at  mileposts  33.1,  33.9,  43.5,  64.8,  89.4,  98.8,  and  117.1;  Aug.  11 
and  13,  1950. 

Streams  on  road  from  Anchorage  to  Potter  2.9  (Campbell  Creek),  10.3,  and  10.6 
miles  from  the  city  limits  of  Anchorage,  Aug.  12,  1950. 

Streams  on  Richardson  Highway  (section  between  Valdez  and  Glenallen)  at 
mileposts  20.5,  25.5  (22.4°  C),  26.3  (see  pi.  7),  29.0,  37.3,  54.7,  62.6  (see  pi.  7), 
81.0  (Squirrel  Creek),  and  87.5  (Rock  Creek),  and  lake  at  milepost  27.3  (22.1°  C), 
Aug.  15,  1950. 

Polycelis  borealis  was  also  identified  in  samples  of  triclads  collected 
by  Dr.  Reece  I.  Sailer  in  July  and  September  1948.  The  following 
localities  were  represented:  Richardson  Highway,  mileposts  8.3,  187.5, 
192.9,  209.7,  223.6  and  239.9,  and  Glenn  Highway,  milepost  117.1. 


174  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  103 

Family  Dendrocoelidae 
Genus  Dendrocoelopsis  Kenk 

Dendrocoelopsis  piriformis,  new  species 

Figure  23;  Plate  6,  Figure  3 

Description. — Sexually  mature  specimens  of  this  rather  broad  and 
plump  species  measure  up  to  15  mm.  in  length  and  3  mm.  in  width. 
In  quietly  gliding  animals,  the  head  is  truncated,  with  a  convex 
frontal  margin  (grasping  organ).  There  is  a  visible  subterminal 
depression  corresponding  to  the  adhesive  surface  of  the  grasping 
organ  or  sucker.  The  rounded  lateral  corners  of  the  head  are  formed 
by  the  auricles,  which  protrude  only  little  laterally,  thus  causing  an 
insignificant  narrowing  (neck)  to  appear  behind  them.  Behind  the 
head,  the  body  margins  widen  gradually.  The  greatest  width  is 
reached  at  the  level  of  the  mouth.  Behind  that  level,  the  body  tapers 
again  to  a  bluntly  pointed  posterior  end.  When  the  animal  is  at 
rest,  the  body  appears  short  and  wide,  about  pear-shaped,  the  body 
margins  often  forming  a  wavy  or  irregular  contour.  Yoimg  specimens 
are  more  slender  and  their  lateral  margins  are  more  nearly  parallel 
when  they  are  in  gliding  motion.  At  rest,  however,  they  assume  the 
same  pyriform  shape  as  the  adults.  In  short,  the  habit  of  the  species 
may  be  described  as  resembling  that  of  European  and  Asiatic  repre- 
sentatives of  the  genus  Bdellocephala  Man,  with  which  it  has  in  com- 
mon the  broad  shape  and  the  anterior  grasping  organ. 

The  two  eyes  are  situated  on  the  dorsal  side  of  the  head.  Their 
distance  from  each  other  amounts  to  somewhat  more  than  one-third 
the  width  of  the  head  at  the  level  of  the  eyes.  The  distance  of  each 
eye  from  the  lateral  margin  is  smaller  than  the  distance  from  the 
frontal  margin. 

The  color  of  the  dorsal  side  is  usually  a  cloudy  brown  or  dark 
brownish  gray.  In  some  specimens,  the  pigment  is  arranged  in 
definite  longitudinal  stripes:  one  sharply  marked  median  stripe  flanked 
on  each  side  by  a  light  (usually  yellow)  band ;  laterad  to  this  band  the 
body  darkens,  losing  its  pigment  again  near  the  lateral  margin.  In 
other  specimens,  lacking  the  two  light  dorsal  bands,  the  back  may  be 
either  uniformly  pigmented  or  may  show  a  more  or  less  distinct  darker 
band  along  the  midline.  Young  specimens  and  striped  adults  clearly 
show  the  finer  disposition  of  the  pigment  in  small  rounded  dots,  each 
dot  apparently  corresponding  to  an  individual  pigment  cell. 

The  pigment  pattern  of  the  dorsal  side  of  the  head  is  quite  charac- 
teristic. A  very  dark  field  between  the  eyes  extends  anteriorly  to 
both  sides  of  the  grasping  organ.  The  area  above  the  organ  is  unpig- 
mented  and  white,  and  so  are  the  two  elongated  ocular  areas  antero- 


FRESH-WATER    TRICLADS    OF    ALASKA — KENT  175 

laterad  to  the  eyes.  A  pair  of  indistinct,  converging  streaks  of  lighter 
coloration  may  run  posteriorly  from  the  lateral  angles  of  the  head. 

The  ventral  side  is  light  gray. 

The  pharynx  is  situated,  in  mature  animals,  behind  the  middle  of 
the  body;  the  mouth,  at  the  beginning  of  the  last  third  of  the  body; 
and  the  genital  aperture,  midway  between  the  mouth  and  the  hind  end. 

The  normal  locomotion  of  the  animal  is  quiet  gliding.  When  dis- 
turbed, it  may  attach  itself  to  the  substratum  (apparently  with  the 
marginal  adhesive  zones)  or  move  by  "crawling."  Young  specimens 
do  not  crawl  as  readily  as  do  adult  ones. 

The  grasping  organ,  or  sucker,  in  living  animals,  appears  as  a  well- 
marked  bulge  on  the  frontal  margin  of  the  head,  showing  a  concave 
ventral  depression.  In  preserved  specimens,  the  frontal  margin  is 
generally  curved  ventrally  and  the  site  of  the  organ  forms  a  thick, 
grooved  rim.  Anatomically,  the  organ  consists  of  glandular  and 
muscular  elements.  The  subterminal  adhesive  surface  is  covered  with 
an  epithelium  devoid  of  rhabdites  and  pierced  by  numerous  gland 
ducts  filled  with  a  granular,  eosinophilic  secretion.  The  cell  bodies 
of  the  glands  are  scattered  through  the  mesenchyme  of  the  anterior 
half  of  the  prepharyngeal  region,  particularly  above  the  intestine. 
The  gland  ducts  run  anteriorly  in  dense  bundles  and,  in  their  terminal 
sections,  are  thickly  swollen  with  secretion.  The  muscular  system  of 
the  organ,  which  could  not  be  analyzed  in  detail  on  account  of  the 
density  of  the  glandular  structures,  has  fibers  attached  to  the  adhesive 
surface,  serving  presumably  as  retractors. 

The  rather  short  and  thick  pharynx  is  structurally  typical  of  the 
family  Dendrocoelidae;  its  internal  muscular  zone  consists  of  inter- 
mingled circular  and  longitudinal  fibers. 

Auricular  sense  organs  are  represented  by  two  bands  of  sensory 
epithelium  extending  posteriorly  from  the  sides  of  the  frontal  margin. 
The  cells  of  this  epithelium  are  less  tall  than  are  those  of  the  sur- 
rounding body  epithelum ;  they  lack  rhabdites  and  are  approached  by 
nerve  fibers  from  the  underlying  mesenchyme.  The  location  of  these 
organs  corresponds  to  the  two  light,  longitudinal  streaks  on  the  sides 
of  the  head  seen  in  the  living  animal. 

The  testes  are  of  moderate  size,  numerous,  densely  packed,  and 
occupy  the  dorsal  half  of  the  mesenchyme,  generally  above  the  intes- 
tinal branches.  They  are  arranged  in  two  wide  areas  on  both  sides 
of  the  midline,  extending  from  the  level  of  the  ovaries  close  to  the 
posterior  end. 

The  ovaries,  situated  at  the  level  of  the  second  pair  of  branches  of 
the  anterior  intestinal  trunk,  show  no  structural  peculiarities. 

The  genital  pore  (pg)  leads  immediately  into  two  cavities,  the  male 
atrium  (am)  and,  to  the  right  and  somewhat  posteriorly,  the  duct  of 


176  PROCEEDINGS    OF    THE    NATIONAL    MUSEUM  vol.  103 

the  copiilatory  bursa  (bd).  No  common  atrium  is  present, 
and  both  the  papUla  of  the  penis  and  the  opening  of  the 
oviduct  are  situated  in  the  "male"  atrium.  The  atrium  is  a  conical 
cavity,  wide  anteriorly  and  tapering  toward  the  genital  pore.  It  is 
lined  with  a  cubical  epithelium,  below  which  occurs  a  thin  layer  of 
fine  circular  muscle  fibers  and  a  thicker  layer  of  coarser  longitudinal 
fibers. 

The  penis  consists  of  a  spherical,  muscular  bulb  and  an  elongated 
papilla.  The  penis  bulb  is  differentiated  into  a  wide  peripheral 
muscular  zone  and  a  more  central  parenchymatic  zone  which  contains 
a  cavity,  the  seminal  vesicle  (vs) .  The  vesicle  is  lined  with  a  glandu- 
lar epithelium  and  its  wall  forms  villuslike  projections.  The  two 
vasa  deferentia  (vd)  penetrate  the  penis  bulb  from  the  anterolateral 
sides  and  open  into  the  seminal  vesicle  on  two  prominent  conical 
papillae  projecting  from  the  anterior  wall  of  the  vesicle  a  short  dis- 
tance from  each  other. 

The  papilla  of  the  penis  is  finger-shaped,  tapering  toward  the  tip, 
and  is  highly  muscular.  It  is  covered  with  a  thin  cubical  epithelium. 
Below  the  epithelium  there  is,  in  the  basal  portion  of  the  papilla,  a 
thin  layer  of  longitudinal  muscles,  below  which  lie  a  stronger  circular 
layer  and  a  second  longitudinal  layer.  In  the  distal  part  of  the 
papUla,  the  external  longitudinal  layer  is  lacking.  The  axis  of  the 
papilla  is  pierced  by  the  ejaculatory  duct  (de)  which  leads  from  the 
seminal  vesicle  to  the  tip  of  the  papilla.  The  duct  has  a  cubical 
epithelium  and  a  strong  coat  of  longitudinal  muscle  fibers. 

The  two  oviducts  approach  the  midline  in  the  region  of  the  copula- 
tory  apparatus,  the  right  one  passing  between  the  atrium  and  the 
bursa  stalk,  and  unite  behind  and  above  the  atrium.  The  common 
oviduct  (ode)  proceeds  ventrally,  then  curves  anteriorly,  and  empties 
into  the  terminal  part  of  the  atrium  close  to  the  genital  pore. 

The  copulatory  bm-sa  (b)  is  a  lobed  sac  lined  with  a  tall  glandular 
epithelium.  The  distal  parts  of  the  epithelial  cells  are  filled  with 
fine  eosinophilic  granules.  The  duct  or  stalk  of  the  bursa  (bd)  is  dif- 
ferentiated into  a  narrow  anterior  section  with  a  weak  muscular  coat, 
which  connects  with  the  bursa,  and  a  posterior  wider  section  which 
bends  ventrally  and  opens  at  the  genital  pore.  The  ceUs  of  the 
anterior  section  resemble  those  of  the  bursa  in  having  similar  granular 
inclusions.  Those  of  the  posterior  section  lack  the  inclusions  and  are 
ciliated.  The  muscle  coat  of  the  duct  consists  of  intermingled  longi- 
tudinal and  circular  fibers. 

Taxonomic  position. — I  have  placed  the  species  in  the  genus 
Dendrocoelopsis  established  originally  for  a  European  species,  D. 
spinosipenis  (Kenk).  The  original  definition  of  the  genus  was  based 
on  the  following  characters:  fibers  of  the  inner  muscle  zone  of  the 


FRESH-WATER    TRICLADS    OF   ALASKA — ^KENK 


177 


pharynx  intermingled ;  no  adenodactyl ;  penis  papilla  developed,  penis 
bulb  of  simple  structure;  oviducts  unite  without  embracing  bursa 
stalk;  zone  of  testes  extending  behind  the  level  of  the  copulatory 
organs;  anterior  end  with  subterminal  true  sucker;  eyes  not  numerous 
(Kenk,  1930).  Subsequently  Beauchamp  (1932,  p.  254)  founded  a 
new  genus,  Amyadenium,  with  two  species,  A.  vandeli  Beauchamp  and 
A.  brementi  (Beauchamp),  both  from  the  Pyrenees.  Two  more 
species  were  reported  by  the  same  author  in  later  papers,  A.  chattoni 
Beauchamp  (1949,  p.  60),  again  from  the  Pyrenees,  and  A.  garmieri 
Beauchamp  (1950,  p.  65),  from  central  France.  Beauchamp  recog- 
nized the  close  relation  of  the  new  genus  to  Dendrocoelopsis,  but 


Figure  23. — Dendrocoelopsis  piriformis,  diagram  of  the  copulatory  organs  in  longitudinal 
section,  X  60.  am,  male  atrium;  b,  bursa;  bd,  bursa  stalk;  de,  ejaculatory  duct;  o,  mouth; 
ode,  common  oviduct;  pg,  genital  pore;  vd,  vas  deferens;  vs,  seminal  vesicle. 

separated  it  from  the  latter  on  accoimt  of  the  absence  of  a  highly 
complex  grasping  organ,  or  true  sucker  (i.  e.,  an  adhesive  organ  sepa- 
rated from  the  surrounding  mesenchyme  by  a  muscle  layer) .  Hyman 
(1935)  described  a  dendrocoelid  species  from  Montana  under  the 
name  of  D.  vaginatus.  Again  the  main  characters  of  the  species 
coincide  with  those  of  D.  spinosipenis  with  the  exception  of  the 
grasping  organ,  which  is  of  a  simpler  type.  The  new  Alaskan  species, 
D.  piriformis,  also  falls  clearly  in  the  vicinity  of  the  species  enumer- 
ated, as  does  another  species,  D.  alaskensis,  the  description  of  which 
follows  on  p.  178.  Within  this  group  of  species  there  is  a  gradual 
differentiation  of  the  grasping  organ,  which  is  absent  in  D.  alaskensis, 
present  as  a  moderately  developed  adhesive  organ  in  D.  vaginata, 
D.  piriformis,  A.  brementi,  A.  vandeli,  A.  chattoni,  and  A.  garmieri, 
and  as  a  more  highly  developed  and  muscular  sucker  in  D.  spinosipenis. 
A  similar  wide  variation  of  the  structure  of  the  anterior  grasping  organ 


178  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  ids 

is  seen  in  the  genus  Dendrocoelum  Orsted  where,  however,  true  suckers 
are  not  known.  In  general,  the  taxonomic  vahie  of  adhesive  organs 
in  triclads  appears  to  be  subordinated  to  that  of  other  anatomical 
structures.  I  therefore  tentatively  modify  the  definition  of  the 
genus  Dendrocoelopsis  by  omitting  the  presence  of  a  sucker  as  a 
generic  character,  to  include  the  species  described  as  Dendrocoelopsis 
and  Amyadenium. 

Dendrocoelopsis  pirijormis  differs  from  the  other  members  of  the 
genus  (in  its  wider  extent)  in  several  characters:  It  is  pigmented, 
whereas  the  others  lack  pigment  and  appear  white;  the  presence  of 
two  eyes  differentiates  it  from  the  three  blind  species,  D.  vandeli,  D. 
brementi,  and  D.  garmieri  and  from  the  many-eyed  D.  chattoni;  and 
the  dorsal  position  of  the  testes  separates  it  from  D.  spinosipenis,  D. 
vaginata,  D.  garmieri,  and  D.  alaskensis.  Apart  from  these  most  con- 
spicuous characters,  the  structure  of  the  male  copulatory  organ  of  D. 
piriformis  is  distinctive. 

Holotype.— On  five  slides,  USNM  22334,  Moose  Creek,  on  Glenn 
Highway,  milepost  186,  near  Alaska  Road  Commission  camp,  Aug. 
U.  1950. 

Distribution  and  Ecology. — Dendrocoelopsis  piriformis  is  a  eury- 
thermic  species  and  inhabits  lakes  and  their  outlets  in  the  southern 
part  of  Alaska  and  also  occurs  in  a  lake  near  Umiat. 

Long  Lake,  on  Glenn  Highway,  milepost  85.9,  Aug.  11,  1950,  clear  water, 
17.8°  C.  (near  bank) ;  under  stones,  several  specimens,  two  of  them  mature. 

Lake  on  Glenn  Highway,  milepost  88.1,  Aug.  11,  1950,  water  temperature 
varying  with  depth  (near  bank,  19°  C.) ;  under  stones,  several  specimens,  one 
mature. 

Lake  on  Glenn  Highway,  milepost  23.5  (see  pi.  8),  Aug.  13,  1950,  clear  water, 
23.6°  C.  (near  bank) ;  several  immature  specimens. 

Stream  crossing  Glenn  Highway  at  milepost  147.2,  outlet  of  Snowshoe  Lake, 
Aug.  14,  1950,  moderate  current,  water  somewhat  colored,  17.0°  C.;  one  young 
specimen,  under  a  stone. 

Moose  Creek,  on  Glenn  Highway,  milepost  186,  near  Alaska  Road  Commission 
camp,  Aug.  14,  1950,  fast,  clear  stream,  16.2°  C;  under  stones,  two  mature  speci- 
mens (holotype). 

Pippin  Lake,  on  Richardson  Highway,  milepost  84.4,  Aug.  15,  1950,  shallow 
water  near  bank,  20°  C. ;  under  stones,  many  specimens,  two  of  them  mature. 

Fresh-water  lake  near  Umiat,  collected  by  P.  F.  Scholander,  Aug.  15,  1948; 
18  specimens,  majority  belonging  to  the  plain  form,  1  with  distinct  stripes,  5 
sexually  mature  (USNM  23678). 

Dendrocoelopsis  alaskensis,  new  species 

Figure  24;  Plate  6,  Figure  4 

Description. — I  had  only  limited  material  of  this  species  and  none 
of  the  animals  was  fully  mature.  The  largest  specimen  measured  20 
mm.  in  length  and  4  mm.  in  width. 

The  anterior  end  is  slightly  lobed,  with  a  convex  frontal  margin 


U.  S.    NATIONAL    MUSEUM 


PROCEEDINGS.    VOL.,    103,    PLATE   7 


Top:  Beaver  pond  at  milepost  62.6  of  Richardson  Highway,  x\laska.  Below  the  beaver 
dam,  Polycelis  borealis  under  stones.  Bottom:  Stream  flowing  along  Richardson  High- 
way, Alaska,  at  milepost  26.3.     On  the  undersides  of  stones,  Polycelis  borealis. 


U.   S.    NATIONAL   MUSEUM 


PROCEEDINGS,   VOL.    103.   PLATE  8 


Top:  Stream  crossing  Sieese  Highway,  Alaska,  at  milepost  84.0.  Habitat  of  Pkagocata 
nivea  and  Dendrocoelopsis  alaskensis.  Bottom:  Lake  at  milepost  23.5  of  Glenn  High- 
way, Alaska.     Under  stones  near  bank,  Dendrocoelopsis  piriformis. 


FRESH-WATER    TRICLADS    OF   ALASKA — ^KENK  179 

and  a  pair  of  rounded  auricles  protruding  both  anteriorly  and  later- 
ally. No  distinct  adhesive  or  grasping  organ  is  developed.  There  is 
a  gentle  narrowing  of  the  head  (neck)  behind  the  auricles.  Behind 
the  head,  the  width  of  the  body  gradually  increases  until  the  greatest 
width  is  reached.     The  posterior  end  is  bluntly  pointed. 

The  body  lacks  pigment,  being  white  except  for  the  contents  of 
the  intestine,  which  may  show  through  the  body  wall. 

There  are  two  principal  eyes.  The  distance  between  them  amounts 
to  about  one-third  the  width  of  the  head  at  the  level  of  the  eyes. 
The  distance  of  each  eye  from  the  frontal  margin  is  equal  to,  or 
slightly  larger  than,  the  distance  from  the  lateral  margin.  Additional, 
supernumerary,  eyes  were  seen  in  a  few  individuals  at  short  distances 
either  anterior  or  posterior  to  the  principal  eyes. 

The  species  bears  a  striking  resemblance  to  a  European  species, 
Dendrocoelum  nausicaae  Schmidt  of  the  Balkan  Peninsula.  It  is 
generally  associated  with  another  triclad  inhabiting  the  same  streams 
in  Alaska,  Phagocata  nivea,  from  which  it  may  be  distinguished  by 
its  larger  size  and  by  the  position  of  the  eyes. 

A  distinct  adhesive  organ  is  not  seen  in  living  animals.  In  histologi- 
cal sections,  a  transverse  band,  pierced  by  numerous  openings  of 
eosinophilic  glands,  is  found  below  the  frontal  margin  of  the  head. 
The  nature  of  the  glands  and  the  local  differentiation  of  the  epithelium 
correspond  entirely  to  the  structure  of  the  submarginal  adhesive  zone 
which  is  seen  in  this  species,  as  well  as  in  other  triclads,  bordering  the 
lateral  margins  of  the  body.  The  frontal  adhesive  area  may,  therefore, 
be  interpreted  as  an  extension  of  the  lateral  adhesive  zone.  It  is 
somewhat  wider  than  the  lateral  one  and  has  no  muscular  differen- 
tiations such  as  are  typical  of  true  grasping  organs.  The  continuity 
of  the  two  zones  is  interrupted  by  a  short  gap  on  each  side  of  the 
head  below  the  auricle. 

The  internal  muscle  zone  of  the  pharynx  consists  of  a  layer  of  inter- 
mingled circular  and  longitudinal  fibers.  This  character  places  the 
species  in  the  family  Dendrocoelidae.  The  anterior  intestinal  trunk 
bears  a  fairly  large  number,  21  to  24  pairs,  of  lateral  branches,  or 
diverticula. 

Of  a  total  of  eight  individuals  of  this  species  at  my  disposal,  five 
were  young,  two  showed  primordia  of  genital  structures,  and  only  one 
had  the  principal  parts  of  the  reproductive  system  developed,  though 
not  fully  differentiated  histologically.  The  description  of  the  genital 
organs,  therefore,  must  be  given  with  certain  reservations. 

The  testes  are  predominantly  ventral,  situated  mainly  below  the 
level  of  the  intestinal  diverticula,  occasionally  extending  farther 
dorsally  between  the  diverticula.  No  clearly  recognizable  testes  were 
seen  behind  the  level  of  the  mouth;  they  may,  however,  appear  there 


180  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  io3 

when  the  animal  matures  completely.  The  ovaries  are  situated 
behind  the  third  or  fourth  lateral  branch  of  the  anterior  iatestiual 
trunk. 

The  genital  pore  (pg)  leads  into  an  undivided  cavity,  the  genital 
atrium  (a),  which  is  narrow  at  the  pore  and  expands  anteriorly.  The 
narrow  posterior  part  receives  the  outlet  of  the  copulatory  bursa  and, 
anteriorly  to  it,  the  opening  of  the  common  oviduct. 

The  penis  has  a  fauiy  large,  spherical  bulb  and  a  plump  and  short 
papilla.  The  bulb  contains  a  cavity  with  irregular  outline,  the  seminal 
vesicle  (vs),  into  which  the  two  vasa  deferentia  (vd)  open  separately. 
The  cavity  of  the  bulb  continues  into  the  broad  papUla  and  opens  at 
its  tip.  The  bulb  has  the  usual  coat  of  muscle  fibers  arranged  in 
concentric  layers.  The  papilla,  which  projects  only  little  into  the 
genital  atrium,  has  two  muscle  layers  underlying  the  outer  epithelium, 
a  circular  layer  and  a  longitudinal  one.  No  glandular  structures  are 
differentiated  in  my  specimen. 

The  two  oviducts  unite,  without  embracing  the  bursa  duct,  dorsally 
to  the  genital  atrium.  The  common  oviduct  (ode)  opens  into  the 
posterior,  narrow  part  of  the  atrium  from  the  dorsal  side. 

The  bursa  (b)  is,  in  my  specimen,  a  rather  small  sac  with  a  narrow 
lumen.  The  bursa  duct  (bd)  runs  posteriorly,  above  the  penis,  to  a 
level  behind  the  genital  pore.  There  it  turns  abruptly  toward  the 
ventral  side.  Its  terminal  portion  is  considerably  wider  than  the 
anterior  part  of  the  duct  and  opens  from  the  posterodorsal  side  into 
the  genital  atrium  close  to  the  genital  aperture.  In  full  maturity,  the 
widened  section  of  the  bursa  duct  probably  represents  a  histologically 
distinct  vagina. 

Taxonomic  position. — The  systematic  relations  of  Dendrocoelopsis 
alaskensis  are  discussed  together  with  those  of  the  preceding  species, 
D.  piriformis.  D.  alaskensis  is  distinguished  from  all  other  species 
of  the  genus  by  the  lack  of  a  grasping  organ. 

Holotype.  On  seven  slides,  USNM  22335,  creek  crossing  Elliot 
Highway  at  milepost  31.0,  July  24,  1950. 

Distribution  and  ecology. — Dendrocoelopsis  alaskensis  is  an  inhabitant 
of  cool,  fast  streams  of  the  White  Mountains  and  usually  shares  its 
habitat  with  Phagocata  nivea.  It  is  a  stenothermic  and  rheophUic 
species. 

Clear  spring  and  creek  on  Steese  Highway,  milepost  82.5,  at  the  Alaska  Road 
Commission  camp,  July  21,  1950;  one  immature  specimen,  under  a  stone. 

Stream  crossing  Steese  Highway  at  milepost  84.0  (see  pi.  8),  altitude  2,700 
feet,  July  21,  1950;  two  immature  specimens,  under  stones,  near  liver  bait. 

Creek  crossing  Elliot  Highway  at  milepost  31.0,  July  24,  1950,  water  tempera- 
ture 3.2°  C;  under  stones,  five  specimens,  two  of  them  with  sexual  structures 
Giolotype) . 


FRESH-WATER    TRICLADS    OF   ALASKA — KENK 


181 


I    I         I 

a   ode  pg      bd 

Figure  24. — Dendrocoelopsis  alaskensis,  diagram  of  the  copulatory  organs  in  longitudinal 
section,  X  70.  a,  genital  atrium;  b,  bursa;  bd,  bursa  stalk;  ode,  common  oviduct; 
pg,  genital  pore;  ph,  pharynx  (extended  through  the  mouth);  vd,  vas  deferens; 
vs,  seminal  vesicle. 

Zoogeographic  conclusions 

The  occurrence  in  Alaska  of  four  endemic  species  of  triclads  poses 
several  interesting  questions.  It  is  known  that  fresh-water  triclads 
are  most  abundantly  represented  in  the  northern  Temperate  Zone 
and  that  the  number  of  species  declines  toward  both  the  Arctic  and 
the  Tropical  Zones.  The  paucity  of  the  triclad  fauna  at  high  lati- 
tudes has  frequently  been  attributed  to  the  effects  of  glaciation  in 
rather  recent  geologic  time.  During  the  glacial  period,  when  huge 
ice  masses  covered  great  parts  of  the  northern  hemisphere,  all  fresh- 
water life  over  wide  areas  must  have  vanished.  After  the  glaciers 
had  receded,  the  areas  were  gradually  repopulated  by  species  entering 
them  from  adjacent  territories.  The  pattern  of  distribution  of 
freshwater  triclads  m  Europe  shows  evidence  of  a  definite  succession 
of  species  which  migrated  into  the  previously  glaciated  areas  in  the 
postglacial  period  (cf.  Thienemann's  1950  summary  of  the  pertinent 
literature) . 

It  is  well  known,  however,  that  the  greater  part  of  Alaska  was 
not  covered  with  ice  in  the  glacial  period.  Geologic  evidence  of 
glaciation  has  been  found  only  in  the  Brooks  Range,  the  Alaska 
Range  (and  areas  south  of  it),  and  in  small  isolated  spots  in  the 
Yukon  and  Kuskokwim  River  Valleys  while  the  remaining  surface 
of  Alaska  remained  free  of  ice  (cf.  Geological  Society  of  America, 


182  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vou  i03 

1945).  We  are  thus  justified  in  assuming  that  the  fresh-water  life 
of  Alaska  was  not  completely  destroyed  even  at  the  peaks  of  glacia- 
tion  when  practically  all  Canada  and  a  considerable  part  of  the 
United  States  were  covered  with  ice  caps. 

The  Alaskan  fresh-water  triclads  show  no  close  relationships  with 
the  present  North  American  triclad  fauna  inhabiting  the  midwestem 
plains  and  the  eastern  and  southern  areas  of  the  continent.  A 
comparison  with  the  fauna  of  the  Rocky  Mountains  is  not  possible 
at  present,  since  the  West  of  the  United  States  and  of  Canada  is 
almost  unexplored  with  regard  to  triclads  and  to  lower  aquatic  in- 
vertebrates in  general.  In  any  event  it  appears  highly  improbable 
that  Alaska  was  populated  by  species  migrating  to  it  from  the  south 
or  east. 

On  the  other  hand,  the  relations  of  the  Alaska  triclads  to  the 
fauna  of  Eurasia  are  unmistakable.  Though,  according  to  our 
present  knowledge,  none  of  the  Alaskan  triclads  is  specifically  identi- 
cal with  any  Asiatic  form,  two  species,  Phagocata  nivea  and  Polycelis 
horealis,  are  very  closely  related  to  Asiatic  species  of  the  same  genera. 

Figure  25  shows  the  geographic  range  of  the  genus  Polycelis  as  it 
is  known  at  present.  The  individual  dots  on  the  map  represent 
either  single  records  or  groups  of  neighboring  localities  where  species 
of  the  genus  have  been  found.  In  interpreting  the  map  it  is  to  be 
kept  in  mind  that  not  all  geographical  areas  are  equally  well  mvesti- 
gated  with  regard  to  the  occurrence  of  lower  invertebrates.  Europe, 
Japan,  and  the  eastern  parts  of  the  United  States  are  comparatively 
well  known  while  our  knowledge  of  the  Asiatic,  Western  American, 
and  African  triclads  is  still  rather  deficient.  A  study  of  the  map 
suggests  that  Polycelis  is  primarily  a  Eurasian  genus.  It  appears 
to  have  extended  its  range,  in  some  earlier  geological  period,  to  the 
northern  rim  of  Africa.*  In  a  similar  way  it  may  have  migrated  into 
Alaska  at  the  time  of  the  cenozoic  land  bridges  and  may  have  pene- 
trated south  along  the  Rocky  Mountains.  It  is  highly  probable 
that  Polycelis  will  be  found,  in  the  future,  more  widely  distributed 
in  the  Rockies  than  present  collection  records  indicate. 

The  repeated  emergence  of  a  land  bridge  between  Alaska  and  the 
eastern  tip  of  Siberia  is  generally  accepted  by  geologists  and  paleon- 
tologists. There  is  ample  evidence  of  an  exchange  of  faunal  elements 
between  the  two  continents  (cf.  Simpson,  1940  and  1947).  The  most 
recent  corridor  must  have  existed  during  the  glacial  stages  of  the 
Pleistocene.  The  volume  of  ice  masses  accumulated  over  vast  areas 
of  the  northern  hemisphere  has  been  estimated  conservatively  at 
34  to  42  million  cubic  kilometers.     The  corresponding  depletion  of 

♦  The  presence  of  Polycelis  oculi-marginata  (Palombi)  in  New  Guinea  is  a  zoogeographic  enigma  (cf.  Beau- 
champ,  1947). 


FRESH-WATER    TRICLADS    OF   ALASKA — KENK 


183 


fe 


184  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  103 

the  ocean  must  have  lowered  the  sea  level  50  to  90  meters  below  the 
present  level  (Daly,  1934,  pp.  41-50).  The  depth  of  the  eastern  part 
of  the  Bering  Sea  (withm  the  peruneter  passing  through  the  eastern 
projection  of  Siberia,  the  Seward  Peninsula  of  Alaska,  and  Nunivak 
and  St.  Lawrence  Islands)  averages  less  than  40  meters.  Thus  the 
ocean  bottom  between  Alaska  and  Asia  must  have  emerged  with  each 
glacial  stage,  forming  a  wide  connection  between  the  two  continents. 
This  process  may  have  been  enhanced  by  the  uplifting  of  the  area 
along  the  margin  of  the  ice  cap,  brought  about  by  plastic  or  elastic 
deformation  of  the  earth  crust  under  the  weight  of  the  ice  masses. 

A  land  bridge  between  Alaska  and  Asia  must  have  persisted  for 
prolonged  periods  of  time.  Simultaneously,  Alaska  was  discon- 
nected from  other  inhabitable  areas  of  North  America  by  broad  ice 
fields. 

In  view  of  the  geological  history  of  Alaska  we  may  conclude  that 
the  present  Alaskan  fresh-water  triclads  are  the  remnants  or  successors 
of  triclads  that  lived  in  Alaska  in  the  glacial  times  and  perhaps  even 
in  preglacial  periods,  and  that  they  are,  in  all  probability,  of  Asiatic 
origin.  It  may  further  be  assumed  that  some  Alaskan  triclads  have 
extended  their  range  during  the  postglacial  period  and  proceeded 
southeast  along  the  Rocky  Mountains.^  The  occurrence  of  a  species 
of  Polycelis  in  Wyoming  and  South  Dakota  may  have  resulted  from 
this  migration.  More  light  would  be  thrown  on  this  question  by  a 
more  thorough  study  of  the  triclad  fauna  of  the  Canadian  and 
American  Rockies. 

Literature  cited 

Arndt,  Waltheb 

1922.  Untersuchungen  an   Bachtricladen.     Ein   Beitrag  zur  Kenntnis  der 

Paludicolen  Korsikas,  Rumaniens  und  Sibiriens.     Zeitschr.  fur  wiss. 

Zool.,  vol.  120,  pp.  98-146,  pi.  4. 
Bazikalova,  a.  fA. 

1947.  Turbellaria-Triclada  vostochnol  Sibiri  i  Pribalkal'fa.     Doklady  Akad. 

Nauk  SSSR,  new  ser.,  vol.  55,  pp.  671-672. 
Beauchamp,  Paul  be 

1932.  Biospeologica.     LVI.     Turbellaries,     Hirudinees,     Branchiobdellidfe. 

Deuxifeme  s^rie.     Arch.  Zool.  Exp^r.  et  Gen.,  vol.  73,  pp.  113-380, 

pis.  6-8. 
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Ceskoslov.  Zool.  Spolecnosti  v  Praze,  vol.  6-7,  pp.  91-96. 
1947.  Observations  sur  quelques  Turbellaries  du   Mus6e  royal  d'Histoire 

naturelle  de  Belgique.     Bull.  Mus.  Roy.  Hist.  Nat.  Belgique,  vol. 

23,  No.  33,  11  pp. 

1949.  Biospeologica.     LXIX.     Turbellaries   (troisifeme  s^rie).     Arch.   Zool. 

Exp^r.  et  G6n.,  vol.  86,  Notes  et  Revue,  pp.  50-65. 

1950.  Nouvelles  diagnoses  de  triclades  obscuricoles.     Bull.  Soc.  Zool.  France, 

vol.  75,  pp.  65-70. 

•  I  wish  here  to  correct  my  assumption  in  a  previous  paper  (Kent,  1943,  p.  6)  that  all  Canadian  triclads 
had  entered  Canada  from  the  south. 


FRESH-WATER    TRICLADS    OF   ALASKA — ^KENK  185 

Bergendal,  D. 

1890.  Studien   iiber  nordische  Turbellarien   und   Nemertinen.     Ofv.    Vet.- 
Akad.  Forh.,  1890,  pp.  323-328. 
Daly,  Reginald  Aldworth 

1934.  The  changing  world  of  the  ice  age.     xxi  +  271  pp.,  6  pis.  New  Haven. 
Geological  Society  of  America 

1945.  Glacial  map  of  North  America.     Special  Papers,  No.  60. 
Girard,  Charles 

1894.  Recherches  sur  les  Planari^s  et  les  N^mertines  de  I'Amerique  du  Nord. 
Ann.  Sci.  Nat.,  ser.  7  (Zool.),  vol.  15,  pp.  145-310,  pis.  3-6. 
Hyman,  Libbie  H. 

1931.  Studies  on  the  morphology,  taxonomy,  and  distribution  of  North 
American  triclad  Turbellaria.  III.  On  Polycelis  coronata  (Girard). 
Trans.  Amer.  Micr.  Soc,  vol.  50,  pp.  124-135. 

1934.  Report  on  triclad  Turbellaria  from  Indian  Tibet.     Mem.  Connecticut 

Acad.  Arts  Sci.,  vol.  10,  No.  2,  pp.  5-12,  pis.  1-2. 

1935.  Studies  on  the  morphology,  taxonomy,   and  distribution  of  North 

American    triclad    Turbellaria.     VI.     A    new    dendrocoelid    from 

Montana,    Dendrocoelopsis   vaginatus   n.   sp.     Trans.  Amer.    Micr. 

Soc,  vol.  54,  pp.  338-345. 
1937.  Studies  on  the  morphology,  taxonomy,   and  distribution  of  North 

American    triclad    Turbellaria.     VII.     The   two   species    confused 

under  the  name  Phagocata  gracilis,  the  validity  of  the  generic  name 

Phagocata  Leidy  1847,  and  its  priority  over  Fonticola  Komdrek  1926. 

Trans.  Amer.  Micr.  Soc,  vol.  56,  pp.  298-310. 
Ijima,  I.,  and  Kaburaki,  T. 

1916.  Preliminary   descriptions   of  some   Japanese  triclads.     Annot.   Zool. 

Japon.,  vol.  9,  pp.  153-171. 
Kaburaki,  Tokio 

1922.  On  some  Japanese  freshwater  triclads;  with  a  note  on  the  parallelism 

in  their  distribution  in  Europe  and  Japan.  Journ.  Coll.  Sci.,  Imp. 

Univ.  Tokyo,  vol.  44,  No.  2,  71  pp.,  1  pi. 
Kenk,  Roman 

1930.  Beitrage  zum  System  der  Probursalier  (Tricladida  paludicola).  III. 

Zool.  Anz.,  vol.  89,  pp.  289-302. 

1936.  Bemerkung  zur  Trikladenfauna  Turkestans.     Zool.   Anz.,   vol.    115, 

pp.  76-80. 
1943.  Notes  on   the   planarian   fauna   of    Canada.     Canadian   Field-Nat., 
vol  57,  pp.  5-6. 
LivANov,  N.  A.,  and  Zabusova,  Z.  I. 

1940.  Planarii   basselna   Teletskogo   ozera  i   novye   dannye   o   nekotorykh 
drugikh  sibirskikh  vidakh  (Paludicoles  du  Lac  Teletzkoe  et  nouvelles 
donnees  sur  quelques  formes  sib^riennes) .     Trudy   Obshch.   Estes- 
tvoisp.  pri  Kazan.  Univ.,  vol.  56,  pp.  83-159. 
MuTH,  Anton 

1912.  Beitrage  zur  Kenntnis  der  Gattung  Sorocelis  Grube.     Mitt.  Naturw. 
Ver.  Steiermark,  vol.  48,  pp.  381-410. 
Palombi,  a. 

1931.  Turbellari    della    Nuova    Guinea.     In:  R^sultats    scientifiques    du 

voyage  aux  Indes  Orientales  Neerlandaises  de  LL.  AA.  RR.  le 
Prince  et  la  Princesse  Leopold  de  Belgique,  vol.  2,  No.  8,  14  pp., 
1  pi.  (M6m.  Mus.  Roy.  Hist.  Nat.  Belgique,  hors  s6ne). 


186  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vou  103 

Sabxjssowa,  Z.  (see  also  Zabusova,  Z.  I.) 

1929.  Die  Turbellarien  der  Kamtschatka-Halbinsel  nach  den  Sammlungen 

der    Rjabuschinsky-Expedition     1908-1909.     Zool.     Jahrb.,     Abt. 

Syst.,  vol.  57,  pp.  497-536,  pi.  4. 
Seidl,  Heinrich  H. 

1911.  Beitrage   zur   Kenntnis   centralasiatischer  Tricladen.     Zeitschr.     ftir 

wiss.  Zool.,  vol.  98,  pp.  31-67,  pis.  5-7. 
Simpson,  George  Gaylord 

1940.  Mammals  and   land   bridges.     Journ.    Washington   Acad.   Sci.,   vol. 

30,  pp.  137-163. 
1947.  Holarctic   mammalian  faunas   and   continental   relationships   during 

the  Cenozoic.     Bull.  Geol.  Soc.  Amer.,  vol  58,  pp.  613-687. 
Thienemann,  August 

1950.  Verbreitungsgeschichte     der    Siisswassertierwelt     Europas;     Versuch 

einer  historischen  Tiergeographie  der  europaischen  Binnengewasser. 

XVI +  809  pp.  Stuttgart. 
Zabusov",  I.  P.  (Sabussow,  H.;  Zaboussoff,  H.) 

1901.  Zamietki  po  morfologii  i  sistematikie  Triclada.     II.     O  planarlfakh" 

Onezhskago     ozera     (Tricladenstudien.     II.     Zur     Kenntniss    der 

Tricladen  des  Onegasees).     Protok.    Zasfed.    Obshch.  Estestvoisp. 

Imp.  Kazan.  Univ.,  Prilozhenle  No.  191,  18  pp. 
1911.  Izsliedovaniia  po   morfologii   i  sistematikie   planarlK   ozera   Balkala. 

I.   Rod"  Sorocelis  Grube   (Untersuchungen  iiber  die  Morphologie 

und  Systematik  der  Planarien  aus  dem  Baikalsee.     I.     Die  Gattung 

Sorocelis    Grube).    Trudy  Obshch.  Estestvoisp.  Imp.  Kazan.  Univ., 

vol.  43,  No.  4,  422  +  8  +  2  pp.,  11  pis. 
1916.  Rjabuschinskya  schmidti  n.  g.  n.  sp.,  novyl  vid"  i  rod"  Tricladida 

paludicola  iz"   Kamchatki   {Rjabuschinskya  schmidti  n.   g.  n.  sp., 

espece  et  genre  nouveau  des  Tricladida  paludicola  du  Kamtchatka). 

Russ.  Zool.  Zhur.,  vol.  1916,  pp.  273-286. 
Zabusova,  Z.  I.  (Saboussoff,  Z.  ;  see  also  Sabussowa,  Z.) 

1936.  Planarii    Kamchatki    (Les    planaires    paludicoles    du    Kamtchatka). 

Kazan.  Gosud.  Univ.,  Uchenye  Zapiski,  Zool.,  vol.  96,  No.  7,  pp. 

141-174. 


U.  S.  60VERNHENT  PRINTING  OFFICEi  Ifll 


PROCEEDINGS  OF   THE   UNITED   STATES  NATIONAL    MUSEUM 


issued 


SMITHSONIAN  INSTITUTION 
U.  S.  NATIONAL  MUSEUM 


Vol.   103  Wajhington:  1953  No.  3323 

REVIEW  OF  THE  INDO-PACIFIC  ANEMONE  FISHES,  GENUS 
AMPHIPRION,  WITH  DESCRIPTIONS  OF  TWO  NEW 
SPECIES. 


By  Leonard  P.  Schultz 


During  the  summer  of  1950  Dr.  Arthur  D.  Welander,  School  o 
Fisheries,  University  of  Washington,  and  I  were  engaged  in  studying 
reef  fishes  brought  back  from  the  Marshall  Islands  by  the  staff  of  the 
Applied  Fisheries  Laboratory  of  the  University  of  Washington. 
Among  this  material  was  a  specimen  of  anemone  fish  that  we  could 
not  identify  with  any  known  species.  After  I  returned  to  the  U.  S. 
National  Museum,  I  reviewed  the  descriptions  of  all  known  species, 
compared  that  specimen  with  the  numerous  lots  of  Amphiprion  in 
the  National  Museum,  and  found  that  it  represented  a  new  species. 
During  March  1951,  I  studied  the  anemone  fishes  in  the  Museum 
of  Comparative  Zoology,  Harvard  University,  and  found  another 
undescribed  species  from  Mauritius. 

Descriptions  and  analyses  of  species  referable  to  the  genus  Amphi- 
prion have  been  based  on  so  few  specimens,  usually  only  one  or  two, 
that  the  problem  of  variability  or  constancy  of  the  color  pattern  has 
been  neglected.  For  most  of  the  few  hundred  species  among  more 
than  fifty  fish  families  that  I  have  studied  in  detail  the  basic  color 
pattern  has  been  observed  to  be  fairly  constant.  It  is  of  great  value 
for  the  recognition  of  species,  especially  in  the  genus  Amphiprion. 
Weber  and  de  Beaufort  (The  fishes  of  the  Indo-Australian  Archipelago, 
vol.  8,  pp.  330-348,  1940)  recognized  eight  species,  whereas  we  have 
distinguished  fom-teen  and  there  may  be  others  recognizable  when 
larger  series  are  compared  and  additional  characters  studied.  Fin 
ray  counts  were  made  on  various  species  and  these  data  are  recorded 
in  table  1. 

Except  for  original  descriptions,  no  attempt  was  made  to  include 
all  references  to  species  referable  to  the  genus  Amphiprion.     Whenever 

238537—53  187 


188  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  io3 

figures  of  species  were  found  these  have  been  mcluded  in  the  synonymy. 
Most  descriptions  and  records  are  not  in  sufficient  detail  to  assign 
them  to  the  correct  species  without  examining  the  material  on  which 
the  records  were  based. 

Genus  Amphiprion  Bloch  and  Schneider 

Amphiprion   Bloch  and   Schneider,   Systema  ichthyologiae  .  .  .,   p.   200,    1801 

(genotype,  Lutjanus  ephippium  Bloch). 
Prochilus   (on   Klein,    1775)    Bleeker,   Nat.   Verh.   Hollandsche   Maatsch,   Wet. 

Haarlem,  ser.  3,  vol.  2,  No.  6,  p.  20,  1877  (genotjrpe,  Lutjanus  ephippium 

Bloch). 
Actinicola  Fowler,  Journ.  Acad.  Nat.  Sci,  Philadelphia,  ser.  2,  vol.  12,  p.  533, 

1904  (genotype,  Lutjanus  percula  Lacep&de). 
Phalerebus  Whitley,  Mem.  Queensland  Mus.,  vol.  9,  pt.  3,  p.  216,  1929  (genotype, 

Prochilus  akallopisos  Bleeker). 

Key  to  the  species  of  Amphiprion 

la.  A  white  band  (sometimes  indistinct)  along  middorsal  line  from  snout  to 
dorsal  origin  or  beyond  along  base  of  dorsal  fin;  total  pectoral  rays  17  to 
19;  next  to  last  dorsal  spine  about  1.3  to  1.5  in  longest  dorsal  spine;  no 
notable  emargination  in  dorsal  fin;  scales  on  dorsal  surface  of  head  extend 
forward  to  a  Une  between  front  of  orbits. 
2a.  No  vertical  pale  bars;  dorsal  rays  about  X,18  or  19;  anal  about  11,12  or 

13;  pectoral  17  to  19 A.  akalJopisos  Bleeker 

26.  A  single  vertical  pale  bar  about  2  scales  wide  from  nape  to  subopercle; 
dorsal    rays    about    X,16;    anal    about    11,12    or    13;    pectoral    17. 

A.  perideraion  Bleeker 
16.   No  white  band  along  middorsal  line. 

3a.  Caudal  fin  with  pale  or  dusky  roundish  center  posteriorly  edged  with 
black;  outer  edges  of  caudal  fin  white;  second  pale  or  white  bar  from 
rear  of  spiny  dorsal  fin  to  anus  with  a  triangular  anterior  projection 
under  depressed  pectoral  fin;  tips  of  pelvics  black;  a  broad  white 
color  bar  on  head  and  on  caudal  peduncle  always  present.  Dorsal 
fin  deeply  indented  at  rear  of  spiny  part,  next  to  last  dorsal  spine 
contained  three  times  in  longest  dorsal  spine;  scales  on  dorsal  surface 

of  head  do  not  extend  forward  of  nape A.  percula  (Lacepede) 

36.  Color  not  as  in  A.  percula. 

4a.  Central  part  of  caudal  fin  black;  outer  edges  of  caudal  fin  broadly  or 

narrowly  edged  with  white;  second  pale  bar,  if  present,  without 

any  projection  anteriorly;  pale  bar  on  head  present;  next  to  last 

dorsal  spine  contained  1.2  to  2.0  times  in  longest  dorsal  spine;  no 

notable  emargination  in  dorsal  fin. 

5o.  Second  pale  bar  represented  dorsally  on  body  by  an  ovate  white 

area  that  continues  anterodistally  on  soft  dorsal  fin;  this  white 

area  does  not  extend  below  midlengthwise  axis  of  body  and 

never  to  anus;  first  pale  bar  10  to  12  scales  wide;  anal  fin  black, 

except  distally  edged  with  white;  spiny  dorsal  black;  pelvics 

black;  pectoral  pale,  except  basally  blackish;  next  to  last  dorsal 

spine  contained  about   1.2  to   1.5  in  longest  dorsal  spine;  no 

notable  emargination  in  dorsal  fin;  scales  on  dorsal  surface  of 

head  extend  forward  to  a  line  between  rear  of  orbits. 

A.  laticlavius  Cuvier  and  Valenciennes 


INDO-PACIFIC    FISH    GENUS    AMPHIPRION — SCHULTZ  189 

56.  Second  pale  bar  continuous  from  dorsal  part  of  body  to  region 
of  anus. 
6a.  Second  pale  bar  broad,  about  7  to  14  scales  wide  at  level  of 
lateral  line  (its  width  there  contained  2  or  less  times  in  width 
of  third  black  bar  at  level  of  lateral  line)  from  whence  it  con- 
tinues posterodorsally  on  soft  dorsal  fin;  pelvics  dusky  to 
blackish;  spiny  dorsal  blackish;  anal  blackish,  at  least  basally; 
scales   on  dorsal   surface  of  head  extend  forward  to  a   line 
between  rear  of  orbits. 
7a.  Caudal  peduncle  with  broad  white  bar;  pectoral  fin  pale  dis- 
tally,  basally  dusky, 

A.  chrysogaster  Cuvier  and  Valenciennes 

76.  Caudal  peduncle  black;   no   white   bar;   black  coloration   of 

posterior  part  of  body  continues  on  central  part  of  caudal 

fin;  pectoral  fin  dusky  in  basal  third.A.  poiymnus  (Linnaeus) 

66.  Second  pale  bar  narrow,  about  2  to  6  scales  wide  at  level  of 

lateral  line,  its  width  there  contained  3.5  or  more  times  in 

width  of  third  black  bar  at  level  of  lateral  line;  caudal  fin  black, 

narrowly  edged  with  white;  anal  fin  pale  to  blackish;  scales 

on  dorsal  surface  of  head  extend  forward  to  over  rear  half  of 

pupil;  pelvics  and  pectorals  pale;  soft  dorsal  black,  edged  with 

white. 

8a.  Second  and  third  (peduncular)  pale  bars  about  2  or  3  scales 

wide,  their  width  at  level  of  lateral  line  contained  7  to 

10  times  in  width  of  third  black  bar  at  level  of  lateral 

line;  second   pale   bar  not  extending  to   distal  edge  of 

dorsal   fin,   ending   on   basal   half   of   last   dorsal   spine; 

posterodorsal  part  of  body  blackish;  anterior  and  ventral 

parts  of  body  pale;  spiny  dorsal  light  dusky. 

A.  tricinctus  Schultz  and  Welander,  new  species 
86.  Second  and  third  (peduncular)  pale  bars  4  to  6  scales  wide, 
their  widths  at  level  of  lateral  Line  contained  from  4  to 
6.5  times  in  width  of  third  black  bar  at  level  of  lateral 
line;  second  pale  bar  extends  into  distal  half  of  dorsal  but 
not  quite  to  the  edge  of  that  fin;  dorsal  and  posterior  half 
of  body  blackish;  anteroventral  part  of  body  pale;  spiny 
dorsal  dusky  to  blackish. 

A.  mauritiensis  Schultz,  new  species 

4a.  Caudal  fin  plain  pale  or  plain  dusky,  no  black  central  blotch  edged 

with  white  posteriorly;  first  pale  bar  on  head  present. 

9a.  Second  pale  bar  broad,  about  7  to  10  scales  wide  at  level  of  lateral 

line,  and  continuing  to  distal  edge  of  spiny  dorsal  fin,  thence 

posteriorly  along  distal  edge  of  soft  dorsal;   caudal  fin  pale; 

caudal  peduncle  black,  without  the  third  pale  bar,  at  least  on 

adults;  spiny  dorsal,  anal,  and  pelvic  fins  black;  pectoral  fin  pale; 

next  to  last  dorsal  spine  contained  about  1.5  in  longest  dorsal 

spine;  scales  on  dorsal  surface  of  head  extend  forward  to  a  line 

between  rear  of  orbits A.  sebae  Bleeker 

96.  Second  pale  bar,  if  present,  not  continuing  to  distal  edge  of  dorsal 
fin;  next  to  last  dorsal  spine  contained  0.8  to  1.2  in  longest  dorsal 
spine;  scales  on  dorsal  surface  of  head  extend  forward  to  lines 
between  rear  edge  of  orbits  to  center  of  pupil. 
10a.  First  two  pale  bars  typically  present  on  adults. 


190  PROCEEDINGS    OF   THE   NATIONAL   MUSEUM  vol.  io3 

llo.  First  two  pale  bars  broad,  each  about  5  to  7  scales  wide  at 
level  of  lateral  line;  third  (peduncular)  pale  bar  represented 
by  a  white  bar,  posteriorly  edged  with  darkish,  or  the  dark 
body  color  on  anterior  part  of  caudal  peduncle  may  end 
abruptly,  the  white  continuing  on  caudal  fin;  spiny  dorsal 
black,  dusky,  or  pale;  soft  dorsal  and  anal  fins  pale  to  black; 
pelvics  pale  to  black,  sometimes  edged  with  black  anteriorly; 
pectoral  pale;  next  to  last  dorsal  spine  about  1.2  in  longest 

dorsal  spine A.  xanthurus  Cuvier  and  Valenciennes 

116.  First  two  pale  bars  narrow,  second  usually  narrower  than  first, 
the  latter  3  to  4.5  scales  wide  and  the  first  3  to  6  scales  wide, 
at  level  of  lateral  line;  caudal  peduncle  blackish,  without 
third  pale  bar  on  adults,  sometimes  pale  bar  is  present  on 
young;  dark  color  of  caudal  peduncle  gradually  fading  into 
pale  color  of  caudal  fin;  spiny  dorsal  pale  to  dusky,  soft 
dorsal  pale;  anal  fin  pale;  pectoral  pale;  pelvics  pale,  except 
anterior  edge  black;  second  from  last  dorsal  spine  contained 
from  1.0  to  1.2  times  in  longest  dorsal  spine. 

A.  bicinctus  Riippell 
106.  First  pale  bar  usually  present  on  head,  about  4  to  6  scales  wide  at 
level  of  lateral  line;  second  pale  bar  lacking,  except  sometimes 
on  small  young  specimens. 
12a.  Anal  fin  pale  or  partly  pale;  pelvics  with  some  pale  area. 
13a.  Anal  fin  pale,  except  distally  edged  with  fine  black  line; 
pelvics  pale,  except  anterior  edge  black;  breast  pale; 
dorsal  spines  X,  soft  dorsal  rays  usually  17. 

A.  ephippium  (Bloch) 
136.  Anal  fin  pale  or  distally  pale  with  basal  half  (or  less) 
dusky  to  blackish;  pelvics  pale  distally  and  blackish 
ventrally,  or  inner  rays  pale  and  outer  rays  broadly 
blackish,  except  distal  tips,  which  are  pale;  dorsal  spines 
IX  or  X  (more  often  IX  than  X),  soft  dorsal  rays  usually 

17  or  18 A.  frenatus  Brevoort 

126.  Anal  and  pelvic  fins  black,  no  pale  areas  anywhere;  dorsal 
spines  X,  soft  dorsal  rays  usually  17. 

A.  melanopus  Bleeker 

Amphiprion  akaJlopisos  Bleeker 

Plate  9,  Figure  A 

Amphiprion  akallopisos  Bleeker,   Nat.  Tijdschr.   Nederl. -Indie,  vol.   4,  p.  281, 

1853. 
Prochilus  akallopisus  Bleeker,  Atlas  ichthyologique  .  .  .  ,  vol.  9,  pi.  400,  fig.  3, 

1878. 
Phalerebus  Whitley  (new  genus),  Mem.  Queensland  Mus.,  vol.  9,  pt.  3,  p.  216, 

1929  (genotype,  Prochilus  akallopisus  Bleeker  =  A.  akallopisos  Bleeker). 

This  species  is  best  recognized  by  the  presence  of  a  wide  white 
band  from  snout  along  middorsal  line  of  head,  thence  posteriorly 
along  each  side  of  base  of  dorsal  fin,  and  ending  on  dorsal  side  of  caudal 
peduncle;  no  white  bars  are  present.  Three  specimens  were  studied: 
USNM  147130,  from  the  Philippine  Islands;  USNM  82781,  from  the 
Fiji  Islands;  and  MCZ  3308,  from  Sabang  Bay. 


INDO-PACIFIC    FISH    GENUS    AMPHIPRION — SCHULTZ  191 

Amphiprion  perideraion  Bleeker 

Plate  9,  Figuke  B 

Amphiprion  perideraion  Bleeker,   Nat.   Tijdschr.   Nederl-Indie,   vol.  9,  p.   437, 

1855. — Montalban,  Bur.  Sci.  Manila  Monogr.  24,  p.  16,  pi.  4,  fig.  1,  1928 

(Philippine  Islands). 
Prochilus  perideraion  Bleeker,  Atlas  ichthyologique  .  .  .  ,  vol.  9,  pi.  400,  fig.  1, 

1878. 
1  Amphiprion  rosenbergi  Bleeker,  Acta  Soc.  Sci.  Indo-Neerl.,  vol.  6,  p.  16,  1859; 

Atlas  ichthyologique  .  .  .  ,  vol.  9,  pi.  402,  fig.  2,  1878. 

This  species  is  best  recognized  by  the  narrow  white  band  on  the 
middorsal  line  of  head,  beginning  between  front  of  eyes  and  extending 
to  dorsal  origin,  and  a  narrow  white  vertical  bar  on  rear  of  head  be- 
hind eye.  Four  specimens  were  studied:  USNM  141032,  141033, 
and  147129,  from  the  Marshall  Islands  and  Borneo;  and  MCZ  33409, 
from  Amboina. 

Amphiprion  percula  (LacepMe) 

Plate  9,  Figure  C 

Lutjanus  percula  LacepSde,  Histoire  naturelle  des  poissons,  vol.  4,  pp.  194,  239, 
240,  1802  (New  Britain). 

Amphiprion  tunicatus  Cuvier  and  Valenciennes,  Histoire  naturelle  des  poissons, 
vol.  5,  p.  399,  pi.  132,  fig.  2,  1830  (Vanicolo)  .—Lesson,  Voyage  ...  La 
Coquille  .  .  .  ,  Zoologie,  vol.  2,  pt.  1,  p.  192,  pi.  25,  fig.  3,  1830  (Port  Praslin, 
New  Ireland;  Doreh,  New  Guinea). 

Prochilus  percula  Bleeker,  Atlas  ichthyologique  .  .  •  ,  vol.  9,  pi.  400,  fig.  2,  1878. 

Anthias  polymnus  var.  (non  Linnaeus)  Bloch,  Naturgeschichte  der  auslandischen 
Fische,  vol.  6,  p.  103,  pi.  316,  fig.  3,  1792. 

Amphiprion  percula  Giinther,  Journ.  Mus.  Godeffroy,  vol.  15,  Andrew  Garrett's 
Fische  der  Siidsee,  pt.  7,  pi.  124,  fig.  A,  1881  (Samoan  Islands). — Day,  The 
fishes  of  India  .  .  .  ,  vol.  2,  p.  379,  pi.  80,  fig.  4,  1878  (Andamans)  .—Montal- 
ban, Bur.  Sci.  Manila  Monogr.  24,  p.  14,  pi.  2,  fig.  2,  1928  (Philippine  Islands). 

Actinicola  percula  Aoyagi,  Coral  Fishes,  Tokyo,  pi.  37,  fig.  2,  1943  (Kakure- 
Kumanomi);  Biogeographica,  Trans.  Biogeogr.  Soc.  Japan,  vol.  4,  No.  1, 
p.  175,  pi.  9,  fig.  2,  1941  (Japan). 

Amphiprion  bicolor  Castelnau,  Proc.  Zool.  Acclim.  Soc.  Victoria,  p.  92,  1873 
(Port  Darwin). 

Actinicola  bicolor  (Castelnau)  Whitley,  Mem.  Queensland  Mus.,  vol.  9,  pt.  3, 
p.  215,  pi.  27,  fig.  2,  1929  (Port  Darwin). 

This  is  the  most  characteristically  colored  species  in  the  genus 
Amphiprion,  and  is  one  of  the  commonest  seen  associated  with  the 
sea  anemone  Discosoma.  The  second  white  bar  has  a  forward  pro- 
jection under  the  depressed  pectoral  fin.  No  other  species  of  Am- 
phiprion observed  by  me  has  the  forward  edge  of  the  second  dark 
bar  with  a  deep  concavity.  The  centers  of  the  dark  bars  on  the 
sides  may  be  pale  brown  to  blackish;  all  dark  bars  are  black  edged. 
I  have  studied  20  lots,  totaling  35  specimens,  from  the  Philippine, 
Solomon,  Palawan,  Schouten,  and  Morotai  Islands. 


192 


PROCEEDINGS    OF   THE    NATIONAL   MUSEUM 


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north  Australia,  near  Darwin.. 

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frenatus 

Philippine  Islands 

Okinawa  and  Japan 

akallopisos 

perideraion 

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mauritiensis 

tricinctus 

xanthurus 

percxda 

sebae 

INDO-PACiriC    FISH    GENUS   AMPHIPRION — SCHULTZ  193 

Amphiprion  laticlavius  Cuvier  and  Valenciennes 

Plate  9,  Figure  D 

Amphiprion  laticlavius  Cuvier  and  Valenciennes,  Histoire  naturelle  des  poissons, 

vol.  5,  p.  394,  pi.  132,  fig.  1,  1830  (New  Guinea). 
Amphiprion  bifasciatus  (non  Bloch)   Montalban,  Bur.  Sci.  Manila  Monogr.  24, 

p.  15,  pi.  3,  fig.  1,  1928  (Philippine  Islands). 
Amphiprion  bifasciatus  annamensis  Chevy,  Travaux  I'lnst.  Oceanogr.  Indochine, 

Mem.  4,  pt.  1,  poissons,  p.  99,  pi.  39,  1932  (Sud-Annam). 
Prochilus  bifasciatus  (non  Bloch)  Bleeker,  Atlas  ichthyologique  .  .  .  ,  vol.  9,  pi.  400, 

fig.  4,  1878. 
Amphiprion  polymnus  (non  Linnaeus)  Aoyagi,  Coral  Fishes,  Tokyo,  pi.  36,  fig.  2, 

1943    (Toaki-Kumanomi) ;    Biogeographica,    Trans.    Biogeogr.    Soc.    Japan, 

vol.  4,  No.  1,  p.  173,  pi.  12,  fig.  4,  1941  (Japan). 
Amphiprion  unimaculatus  (non  Meuschen)   Okada  and  Ikeda,  Biogeographica, 

Trans.  Biogeogr.  Soc.  Japan,  vol.  3,  No.  2,  p.  202,  fig.  28,  1939  (Itoman, 

Okinawa) . 

This  species  differs  from  all  others  in  regard  to  the  second  white 
bar,  which  is  represented  by  an  ovate  white  area  on  upper  half  of 
body  and  on  soft  dorsal  fin,  but  without  any  extension  on  ventral  part 
of  body.  Third  white  bar  on  caudal  peduncle  is  lacldng.  Caudal 
fin  is  basally  and  centrally  blackish,  with  posterolateral  edges  white. 
Anal  is  submarginally  blackish,  with  distal  edge  white.  One  specimen, 
USNM  147128,  from  the  Philippines,  was  studied. 

Amphiprion  chrysogaster  Cuvier  and  Valenciennes 

Plate  9,  Figure  E 

Amphiprion  chrysogaster  Cuvier  and  Valenciennes,  Histoire  naturelle  des  poissons, 
vol.  5,  p.  400,  1830  (lie  de  France) — Lesson,  Voyage  ...  La  Coquille,  .  .  .  , 
Zoologie,  vol.  2,  pt.  1,  p.  191,  pi.  28,  fig.  3,  1830  (lie  de  France). 

Amphiprion  percula  (non  Lacep&de)  Okada  and  Ikeda,  Biogeographica,  Trans. 
Biogeogr.  Soc.  Japan,  vol.  3,  No.  2,  p.  200,  pi.  6,  fig.  1,  1939  (Riu  Kiu  Islands), 

Amphiprion  trifasciatus  Cuvier  and  Valenciennes,  Histoire  naturelle  des  poissons, 
vol.  5,  p.  695,  1830  (Moluccas). 

Amphiprion  fusciventer  Bennett,  Proc.  Comm.  Zool,  Soc.  London,  vol  1,  p.  165, 
1831  (Mauritius). 

Prochilus  bifasciatus  (non  Bloch)  Bleeker,  Atlas  ichthyologique  .  .  .  ,  vol.  9, 
pi.  400,  fig.  6,  1878. 

This  species,  with  three  broad  white  bars,  has  the  central  part 
of  caudal  fin  black  and  outer  edges  white;  second  white  bar  is  con- 
tinuous on  distal  part  of  soft  dorsal  fin.  Seven  specimens  were 
studied:  One  each  in  USNM  61690  from  Mauritius,  141034  from  the 
Marshall  Islands,  147127  from  the  Philippines;  and  four  MCZ  speci- 
mens, three  from  Zanzibar,  Africa,  and  one,  collected  by  Andrew 
Garrett,  from  Apiang,  Kingsmill  Islands. 


194  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  io3 

Amphiprion  polymnus  (Linnaeus) 

Plate  9,  Figure  J 

Perca  ^olymna  Linnaeus,  Systema  naturae,  ed.  10,  p.  291,  1758. 

Anthias  bifasciatus  Bloch,  Naturgeschichte  der  auslandischen  Fische,  vol.  6,  p. 

103,  pi.  316,  fig.  2,  1792. 
Prochitus  bifasciatus  Bleeker,  Atlas  ichthyologique  .  .  .  ,  vol.  9,  pi.  400,  fig.  5, 

1878. 
? Amphiprion  ocellaris  Cuvier  and  Valenciennes.  Histoire  naturelle  des  poissons, 

vol.  5,  p.  399,  1830  (Sumatra). 
? Amphiprion  melanurus  Cuvier  and  Valenciennes,  Histoire  naturelle  des  poissons, 

vol.  5,  p.  400,  1830  (Sumatra). 
Lutjanus  jourdin  LacepSde,  Histoire  naturelle  des  poissons,  vol.  4,  pp.  191,  235, 

1802  (Amboina). 
Coracinus  seu  Sciaena  unimaculata  Meuschen,  Zoophylacium  Gronovianum  .  .  .  , 

Pisces,  No.  227,  1781  (based  on  Gronow,  1763). 
Coracinus  vittatus  Gray,  Catalogue  of  fish  collected  and  described  by  L.  T.  Gronow 

.  .  .  ,   British  Museum,  p.  57,  1854  (based  on  Gronow's  No.  227). 
Amphiprion  intermedius  Schlegel  and  Miiller,  Verhandelingen  over  de  Natuurlijke 

Geschiedenis  der  Nederlandsche  Overzeesche  Bezittingen  .  .  .  ,  Zoologie, 

p.  18,  1839-1841  (reference  copied). 

Amphiprion  polymnus  (Linnaeus)  has  been  confused  almost  since 
the  day  it  was  named.  Weber  and  de  Beaufort  (Fishes  of  the  Indo- 
AustraUan  Archipelago,  vol.  8,  p.  344,  1940)  discuss  the  confusion 
between  polymnus  and  bicinctus,  the  former  name  having  been  used 
for  the  species  currently  called  bicinctus. 

In  A.  polymnus  the  second  white  bar  continues  on  to  the  distal 
part  of  the  soft  dorsal,  but  it  lacks  the  third  white  bar  on  the  caudal 
peduncle;  also,  the  black  coloration  of  the  caudal  peduncle  continues 
on  to  the  central  part  of  the  caudal  fin,  whereas  in  A.  sebae  the  caudal 
fin  is  pale.    The  anal  fin  is  black  basally,  with  the  distal  third  white. 


Explanation  for  Plate  9 

A,  Amphiprion  akallopisos  Bleeker,  photograph  of  a  color  drawing  in  the  Philippine  Albatross 
collection;  B,  A.  perideraion  Bleeker,  photograph  of  a  color  drawing  in  the  Philippine 
Albatross  collection;  C,  A.  percula  (Lacepede),  photograph  of  a  color  drawing  in  the 
Philippine  Albatross  collection;  D,  A.  laticlavius  Cuvier  and  Valenciennes,  photograph 
of  plate  36,  figure  2,  in  Aoyagi,  Coral  Fishes,  1943  {=  A.  polymnus,  non  Linnaeus,  AoyagI); 
E,  A.  chrysogaster  Cuvier  and  Valenciennes,  photograph  of  plate  6,  figure  1,  In  Okada  and 
Ikeda,  Blogeographica,  Trans.  Blogeogr.  Soc.  Japan,  vol.  3,  No.  2, 1939  {^=A.  percula,  non 
Lacepede,  Okada  and  Ikeda);  F,  Amphiprion  sebae  Bleeker,  photograph  of  figure  27  in 
Okada  and  Ikeda^  Blogeographica,  Trans.  Blogeogr.  Soc.  Japan,  vol.  3,  No.  2,  1939; 
G,  A.  xanthurus  Cuvier  and  Valenciennes,  copy  of  figure  4  in  Jordan  and  Dickerson,  Proc. 
U.  S.  Nat.  Mus.,  vol.  34,  p.  611,  1908,  of  a  specimen  from  Suva,  Fiji;  H,  A.  tricinctus, 
new  species,  holotype,  USNM  152929,  from  Bikini  Atoll,  Amen  Island,  standard  length 
75  mm.;  I,  A.  mauritiensis,  new  species,  holotype,  MCZ  6093,  from  Mauritius,  standard 
length  111.5  mm.;  J,  A.  polymnus  (Linnaeus),  photograph  of  plate  316,  figure  2,  Bloch, 
Naturgeschichte  der  auslandischen  Fische,  vol.  6,  1792  {= Anthias  bifasciatus 'Sioch). 


U.    S.    NATIONAL    MUSEUM 


PROCEEDINGS,    VOL.    103.    PLATE  9 


Certain  species  of  Amphiprion 
(For  explanation,  see  facing  page) 


U.    S.    NATIONAL    MUSEUM 


PROCEEDINGS,    VOL.    103.   PLATE   10 


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INDO-PACIFIC    FISH    GENUS   AMPHIPRION — SCHULTZ  195 

I  have  examined  one  specimen  collected  Jmie  16,  1948,  by  Dr. 
Robert  R.  Miller  in  "a  submerged  canoe  on  a  sand  bar  in  Little 
Lagoon,  northeast  end  of  Groote  Eylandt,  Gulf  of  Carpentaria, 
Australia."  The  color  when  alive  was  as  follows:  "The  pale  bars 
were  coral  pink,  with  narrow  emerald  borders;  the  dark  bars  were 
velvet-black." 

Amphiprion  tricinctus  Schultz  and  Welander,  new  species 

Plate  9,  Figure  H 

Amphiprion  ephippium  (non  Bloch)  var.  chrysopterus  (non  Cuvier  and  Valen- 
ciennes) Giinther,  Journ.  Mus.  Godeffroy,  vol.  15,  Andrew  Garrett's  Fische 
der  Siidsee,  pt.  7,  pp.  224-225,  pi.  122,  fig.  C,  1881  (Kingsmill  Islands). 

Holotype. — USNM  152929,  Bikini  Atoll,  Amen  Island,  lagoon, 
August  21,  1947,  University  of  Washington,  Staff  of  Applied  Fisheries 
Laboratory,  standard  length  75  mm. 

Description. — Dorsal  fin  rays  X,17;  anal  11,14;  pectoral  ii,17,i  to 
ii,16,ii;  pelvics  1,5;  branched  caudal  fin  rays  8  +  7;  vertical  scale  rows 
from  upper  edge  of  gill  opening  to  base  of  caudal  fin  54,  scales  between 
lateral  line  and  base  of  first  soft  dorsal  ray  5,  and  between  lateral 
line  and  anal  origin  20;  pores  in  lateral  line  36;  predorsal  scales  19  or 
20;  gill  rakers  5  +  1  +  13. 

Detailed  measurements  were  made  on  the  holotype  and  these  data 
are  expressed  in  thousandths  of  the  standard  length,  75  mm.:  Greatest 
depth  560;  length  of  head  272;  snout  99;  eye  95;  least  preorbital  35; 
length  from  snout  tip  to  rear  edge  of  maxillary  101 ;  postorbital  length 
of  head  147;  least  width  of  interorbital  space  100;  least  depth  of 
caudal  peduncle  160;  length  of  caudal  peduncle  from  base  of  last  anal 
ray  to  midbase  of  caudal  fin  192;  length  of  longest  ray  of  pectoral  287, 
pelvic  313,  upper  caudal  fin  307,  lower  lobe  of  caudal  fin  300,  spiny 
dorsal  160;  length  of  next  to  last  dorsal  spine  160;  width  of  white 
part  of  first  pale  bar  at  level  of  lateral  line  73,  second  40,  last  (pe- 
duncular) 20. 

Depth  of  body  1.7,  head  3.4,  both  in  standard  length.  Snout  3.2; 
eye  3.1;  least  preorbital  distance  8.0;  upper  jaw  2.8;  postorbital  part 
of  head  2.0;  least  interorbital  space  3.0;  least  depth  of  caudal  peduncle 
1.8;  length  of  pectoral  fin  1.0,  pelvic  0.9,  second  dorsal  spine  23, 
upper  caudal  rays  0.9  to  1.0;  all  in  length  of  the  head.  Least  depth 
of  caudal  peduncle  in  its  length  1.2.  Angle  of  upper  profile  of  head 
with  lengthwise  axis  of  body  about  50° ;  profile  of  head  convex. 

Teeth  in  both  jaws  in  a  single  row,  nearly  conical,  a  little  com- 
pressed forward,  pointed;  interorbital  space  scaled  forward  to  a  line 
between  middle  of  pupils ;  4  or  5  rows  of  scales  on  cheeks ;  gill  cover  with 
a  few  scales;  scales  occur  part  way  out  on  all  median  fins;  preorbital 
with  3  spines;  suborbital  with  10  to  12  smaller  spines. 


196  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  103 

Color  in  alcohol. — Background  coloration  of  body  from  about  fifth 
dorsal  spine  posteriorly  and  dorsally  to  midlengthwise  axis  of  body 
blackish,  the  anteroventral  part  of  body  pale  light  brown  becoming 
paler  ventrally;  spiny  dorsal  dark  brown;  soft  dorsal  black;  caudal  fin 
black,  except  edged  with  white  posteriorly;  pectoral  and  pelvic  fins 
pale  or  very  light  tan ;  anal  pale  or  light  tan,  distally  edged  with  a  black 
line;  first  white  bar  begins  a  little  in  front  of  dorsal  origin  and  just 
behind  eye,  ending  on  lower  edge  of  subopercle;  second  white  bar 
begins  on  last  dorsal  spine  and  base  of  first  soft  ray  in  lower  third  of 
fin,  extends  ventrally,  meeting  its  fellow  in  narrow  space  between 
anal  origin  and  anus;  third  white  bar,  about  half  width  of  second,  or 
narrower  than  width  of  pupil,  crosses  caudal  peduncle  just  in  front  of 
caudal  fin  base. 

Remarks. — This  new  species  may  be  recognized  by  the  narrowness 
of  the  three  white  bars,  especially  the  second  and  third,  and  by  the 
black  caudal  fin  narrowly  edged  with  white  posteriorly.  It  is  sep- 
arated from  all  other  species  of  Amphriprion  by  the  key. 

Named  tricinctus  in  reference  to  the  three  white  bars. 

Amphiprion  mauritiensis  Schultz,  new  species 

Plate  9,  Figure  I 

Holotype. — MCZ  6093,  Mauritius,  collected  by  Nicolas  Pike,  stand- 
ard length  111.5  mm. 

Paratypes. — Bearing  same  data  as  holotype:  Out  of  MCZ  6093,  4 
specimens,  67  to  96  mm.;  MCZ  5801,  2  specimens,  98  and  104.5  mm.; 
MCZ  5800,  1  specimen,  101.5  mm.;  MCZ  5802,  2  specimens,  83.5  and 
114  mm. 

Description. — Dorsal  rays  X,17  or  18,  one  with  XI, 16,  usually 
X,17;  anal  111,14,  one  with  111,13;  pectoral  rays  20  or  21;  pel  vies 
1,5;  branched  caudal  fin  rays  8+7;  vertical  scale  rows  from  upper 
edge  of  gill  opening  to  base  of  caudal  fin  about  55  to  58;  scales  between 
lateral  line  and  base  of  first  soft  dorsal  ray  5 ;  between  lateral  line  and 
anal  origin  17  to  19;  pores  in  lateral  line  35  to  42;  predorsal  scales 
about  10  or  11;  gill  rakers  5  +  1  +  13. 

Detailed  measurements  were  made  on  the  holotype  and  two  para- 
types, and  these  data  are  expressed  in  thousandths  of  the  standard 
length,  first  for  the  holotype  then,  in  parentheses,  for  the  paratypes: 
Standard  length  in  mm.  111.5  (87.5;  104.5).  Greatest  depth  485 
(490;  535);  length  of  head  297  (286;  310);  snout  90  (97;  96);  eye  89 
(86;  86);  least  preorbital  width  31  (38;  28);  distance  from  snout  tip 
to  rear  edge  of  maxillary  91  (114;  112);  least  width  of  interorbital 
space  74  (91;  81);  least  depth  of  caudal  peduncle  144  (157;  151); 
length  of  caudal  peduncle  166  (153;  148).     Length  of  longest  ray  of 


INDO-PACIFIC    FISH    GENUS    AMPHIPRION — SCHULTZ  197 

pectoral  279  (269;  280);  pelvic  291  (257;  296);  upper  caudal  lobe  270 
(280;  296);  lower  caudal  lobe  265  (263;  277);  spiny  dorsal  135  (132; 
124).  Length  of  next  to  last  dorsal  spine  99  (86;  86).  Width  of 
first  white  bar  at  level  of  lateral  line  81  (80;  67);  second  71  (74;  57); 
third  (peduncular)  48  (63;  48).  Width  of  second  black  bar  at  level 
of  lateral  line  323  (331;  344);  third  300  (292;  344). 

Depth  of  body  1.8  to  2.1,  head  3.2  to  3.5,  both  in  standard  length. 
Snout  2.9  to  3.2;  eye  3.3  to  3.6;  least  preorbital  7.5  to  10.1;  upper 
jaw  2.5  to  3.2;  least  interorbital  3.1  to  3.9;  least  depth  of  caudal 
peduncle  1.9  to  2.0,  length  of  pectoral  1.1,  pelvic  1.0  to  1.1,  third 
dorsal  spine  2.1  to  2.5;  all  in  length  of  head.  Least  depth  of  caudal 
peduncle  in  its  length  0.95  to  1.2.  Width  of  second  white  bar  at 
level  of  lateral  line  in  width  of  thu'd  black  bar  at  level  of  lateral  line 
3.5,  thhd  white  bar  4.0  to  7.0. 

Profile  of  head  convex.  Teeth  uniserial,  nearly  conical,  a  little 
compressed  forward,  pointed;  interorbital  space  scaled  forward  to  a 
line  between  rear  of  eyes;  4  to  6  rows  of  scales  on  cheek;  gill  cover 
with  a  few  scales;  scales  occur  part  way  out  on  all  median  fins;  pre- 
orbital with  1  to  3  spines;  suborbital  with  about  a  dozen  spines. 

Color  in  alcohol. — Background  coloration  dark  brownish  or  blackish, 
except  that  underside  of  head  and  ventrally  below  a  line  from  pectorals 
to  anus  is  white  or  pale;  three  white  bars,  one  behind  eye,  second  from 
origin  of  soft  dorsal  to  anus,  third  on  caudal  peduncle;  dorsal  fin 
black,  with  distal  edge  of  soft  dorsal  white;  anal  fin  black  or  pale,  if 
black,  distal  edge  is  white;  pelvics  pale;  pectoral  pale,  except  dusky 
basally  on  some  specimens;  central  area  of  caudal  fin  blackish,  edges 
white,  with  edge  widest  distally;  gill  membranes  appear  to  be  white. 

Remarks. — This  new  species  is  most  closely  related  to  A.  tricinctus 
but  differs  in  having  wider  white  bars,  as  compared  in  the  key. 
Named  mauritiensis  in  reference  to  the  locality  where  it  was  collected. 

Amphiprion  sebae  Bleeker 

Plate  9,  Figure  F 

Amphiprion  sebae  Bleeker,  Nat.  Tijdschr.  Nederl.-Indie,  vol.  4,  p.  478,  1853; 
Atlas  ichthyologique  ,  .  .,  vol.  9,  pi.  400,  fig.  9,  1878. — Day,  Fishes  of 
India  .  .  .  ,  vol.  2,  p.  378,  pi.  80,  fig.  3,  1878  (Andamans) .— Okada  and 
Ikeda,  Biogeographica,  Trans.  Biogeogr.  Soc.  Japan,  vol.  3,  No.  2,  p.  200, 
fig.  27,  1939  (Isigaki  and  Irimote  Islands). 

This  species,  in  which  the  second  white  color  bar  continues  distally 
on  the  soft  dorsal  fin,  is  very  much  like  A.  polymnus,  but  differs  in 
having  a  white  bar  on  the  caudal  peduncle  and  a  white  caudal  fin. 
Three  specimens  were  studied:  USNM  45169,  from  the  Seychelles, 
and  133830,  from  the  Paumotu  Islands;  and  MCZ  33410,  from  Sumatra. 


198  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  los 

Amphiprion  xanthurus  Cuvier  and  Valenciennes 

Plate  9,  Figure  G 

Amphiprion  xanthurus  Cuvier  and  Valenciennes,  Histoire  naturelle  des  poissons, 

vol.  5,  p.  402,  1830  (lie  de  France). 
Amphiprion  clarckii  Cuvier  and  Valenciennes,  Histoire  naturelle  des  poissons, 
ri     vol.  9,  p.  504,  1833  (emended  spelling  on  Bennett). 
Anthias  clarkii  Bennett,  A  selection  from  .  .  .  fishes  found  on  the  coast  of  Ceylon, 

London,  ed.  2,  p.  29,  pi.  29,  1834  (Ceylon). 
Sparus  milli  Bory  de  Saint- Vincent,  Dictionnaire  classique  d'histoire  naturelle, 

vol.  17,  p.  130,  pi.  113,  fig.  2,  1831  (China  Sea). 
Prochilus  polymnus  (non  Linnaeus)  Bleeker,  Atlas  ichthyologique,  vol.  9,  pi.  400, 

figs.  7,  8,  1878. 
Anthias    polymna    (non    Linnaeus)    Bloch,    Naturgeschichte  der  auslandischen 

Fische,  vol.  9,  p.  89,  pi.  316,  fig.  1,  1792. 
Amphiprion  bicinctus  (non  Riippell)  Aoyagi,  Coral  Fishes,  Tokyo,  pi.  37,  fig.  1, 

1943  (Kumanomi). — Biogeographica,  Trans.  Biogeogr.  Soc.  Japan,  vol,  4, 

No.  1,  p.  169,  pi.  9,  fig.  1,  1941  (Japan). 
Amphiprion  polymnus  (non  Linnaeus)  Okada  and  Ikeda,  Biogeographica,  Trans. 

Biogeogr.  Soc.  Japan,  vol.  3,  No.  2,  p.  204,  fig.  30,  1939  (Riu  Kiu  Islands).— 

Montalban,  Bur.  Sci.  Manila  Monogr.  24,  p.  10,  pi.  1,  fig.  1,  1928  (Philippine 

Islands) . 
Amphiprion  chrysopterus  Cuvier  and  Valenciennes,  Histoire  naturelle  des  poissons, 

vol.  5,  p.  401,  1830  (no  locality  given). — Jordan  and  Dickerson,  Proc.  U.  S. 

Nat.  Mus.,  vol.  34,  p.  612,  fig.4,  1908  (Suva,  Fiji). 
Amphiprion  japonicus  Temminck  and  Schlegel,  Fauna  Japonica  .  .  .  ,  Pisces, 

p.  66,  1843  (Japan). 
Amphiprion  chrysargyrus  Richardson,  Rep.  Meetings  British  Assoc.  Adv.  Sci., 

vol.  15  (1845),  p.  254,  1846  (Seas  of  China  and  Japan). 
Amphiprion  boholensis  Cartier,  Verh.  phys.  med.  Wiirzburg,  new  ser.,  vol.  5, 

p.  96,  1874  (Bohol). 
Amphiprion  melanostolus  Richardson,  Ann.  Mag.  Nat.  Hist.,  vol.  9,  p.  390,  1842 

(Depuch  Island). 
7 Amphiprion  de  bojer  Lienard,  TreiziSme  Rapp.  Ann.  Soc.  Hist.  Nat.  Maurice, 

p.  68,  1843  (reference  copied). 
Amphiprion  snyderi  Ishikawa,  Proc.  Nat.  Hist.  Tokyo  Mus.,  vol.  1,  No.  1,  p.  11, 

pi.  5,  1904  (Bonin  Island). 

I  have  examined  70  specimens  (in  45  lots)  of  xanthurus,  22  of  which 
are  in  the  Museum  of  Comparative  Zoology,  Harvard  University, 
and  the  others  in  the  U.  S.  National  Museum,  and  I  find  some  variation 
in  color  pattern.  Among  these  specimens  12  had  black  dorsal,  anal, 
and  pelvic  fins;  3  had  these  three  fins  dusky;  10  had  the  dorsal  and  the 
anal  black  or  dusky,  with  pelvics  pale;  36  had  the  dorsal  black  and 
both  anal  and  pelvics  pale ;  9  had  dorsal,  anal,  and  pelvics  pale,  except 
in  some  specimens  the  pelvic  fins  were  dark  edged.  Of  these  70 
specimens  42  were  from  the  Philippines,  4  were  from  Japan,  2  from 
the  China  coast,  18  from  Zanzibar,  1  from  the  Gilbert  Islands,  and 
3  from  Kingsmill  Islands. 


INDO-PACIFIC    FISH    GENUS   AMPHIPRION — SCHULTZ  199 

Amphiprion  bicinctus   Riippell 

Plate  10,  Figure  A 

Amphiprion  bicinctus  Riippell,  Atlas  zu  der  Reise  im  nordlichen  Airika,  p.  139, 

pi.  35,  fig.  1,  1828  (Red  Sea). 
Amphiprion  papuensis  Macleay,  Proc.  Linn.  Soc.  New  South  Wales,  vol.  8,  No. 

2,  p.  271,  1883  (New  Guinea). — Whitley,  Mem.  Queensland    Mus.,  vol.  9, 

pt.  3,  p.  210,  pi.  27,  fig.  1,  1929  (D'Entrecasteaux  Group,  New  Guinea,  on 

holotype) . 
Amphiprion  arion  De  Vis,  Proc.  Linnean  Soc.  New  South  Wales,  vol.  8,  p.  450, 

1884  (South  Seas). 

This  species  has  the  first  two  white  bars  but  lacks  the  peduncular 
bar;  the  anterior  edges  of  the  pel  vies  are  black  and  the  caudal  fin  is 
pale.  Six  lots  containing  seven  specimens  were  studied:  USNM 
61679,  from  Suva,  Fiji;  USNM  141030  and  141031,  from  the  Marshall 
Islands ;  uncataloged  specimens,  one  from  the  Paumotu  and  two  with- 
out data;  also  one  from  Bikini  in  the  University  of  Washington 
collection. 

Among  these  specimens,  four  adults,  82  to  99  mm.  in  standard  length, 
have  a  pale  anal  fin,  whereas  two,  26  and  27  mm.  long,  have  a  black 
anal,  and  one,  25  mm.  long,  has  a  dusky  anal  fin.  The  caudal  peduncle 
varies  from  pale  dusky  to  black  or  brownish.  The  key  gives  the 
essential  color  pattern  of  this  species. 

Amphiprion  ephippium  (Bloch) 

Plate  10,  Figures  B,  C 

Luijanus  ephippium  Bloch,  Naturgeschichte  der   auslandischen  Fische,  vol.  4, 

p.  121,  1790  (reference  not  seen);  Ichthyologie,  ou  histoire  naturelle,  generate 

et  particulifere  des  poissons  .  . .  ,  vol.  7,  p.  98,  pi.  250,  fig.  2,  1797  (East  Indies). 
Amphiprion  ephippium  Day,  Fishes  of  India  .  .  .  ,  vol.  2,  p.  378,  pi.  80,  fig.  1, 

1878. 
Amphiprion  monofasciatus  Thiolliere  in  Montrouzier,  Suite  de  la  faune  de  I'lle 

de  Woodlark  ou  Moiou,  Ichthyologie,  Ann.  Sci.,  Physic.  Nat.  Agr.  Indust., 

Lyon,  vol.  8,  p.  476,  1856  (Woodlark  Island). 
Amphiprion  tricolor  Gtinther,  Catalogue  of  the  fishes  in  the  British  Museum, 

vol.  4,  p.  8,  1862  (Port  Essington;  South  Australia). 
Amphiprion  riippeli  Castelnau,  Proc.  Zool.  Acclim.  Soc.  Victoria,  vol.  2,  p.  91, 

1873  (Port  Darwin). 
Amphiprion  frenatus  (non  Brevoort)  Day,  Fishes  of  India  .  .  .  ,  vol.  2,  p.  378, 

pi.  80,  fig.  2,  1878. 
Prochilos  ephippium  Bleeker,  Atlas  ichthyologique  .  .  .  ,  vol.  9,  pi.  401,  figs. 

1,  9,  1878. 
Amphiprion  rubrocinctus  Richardson,  Ann.  Mag.  Nat.  Hist.,  vol.  9,  p.  391,  1842 

(Depuch  Island;  probably  young). 

The  adults  of  this  species  usually  have  a  plain  blackish  body  and 
head,  with  a  single  white  color  bar  on  the  head,  but  on  a  few  speci- 
mens, the  largest  in  the  series,  this  first  white  bar  is  lacking  or  nearly 


200  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  los 

SO.  The  smallest  specimens  among  57  collected  by  Dr.  Robert  R. 
Miller  from  Woods  Inlet  west  of  Darwin,  Australia,  have  a  variable 
color  pattern  of  two  or  three  white  bars.  One  specimen,  20  mm.  in 
standard  length,  has  only  one  white  bar  on  the  head;  9,  from  20  to 
31  mm.,  have  two  white  color  bars,  the  first  and  second;  7,  from  19.5 
to  28  mm.,  have  the  first  and  second  white  bars  distinct  and  the  third 
indistinct,  sometimes  represented  by  a  white  spot  on  dorsal  edge  of 
caudal  peduncle;  10,  from  17  to  25  mm.,  have  all  three  white  color 
bars  distinct.  I  presume  that  the  second  and  third  white  bars  dis- 
appear with  increase  in  size,  since  a  35-mm.  specimen  in  this  lot  looks 
like  all  the  larger  specimens  with  only  one  white  bar  on  the  head. 

Amphiprion  frenatus  Brevoort 

Plate  10,  Figure  D 

Amphiprion  frenatus  Brevoort,  U.  S.  Japan  Exped.  Nat.  Hist.,  Washington, 
vol.  2,  p.  263,  pi.  6,  fig.  4,  1856  (Lew  Chew  [Okinawa]). — Jordan  and  Snyder, 
Proc.  U.  S.  Nat.  Mus.,  vol.  24,  p.  597,  1902.— Montalban,  Bur.  Sci.  Manila 
Monogr.  24,  p.  12,  pi.  2,  fig.  1,  1928  (Philippine  Islands). — Aoyagi,  Biogeo- 
graphica.  Trans.  Biogeogr.  Soc.  Japan,  vol.  4,  No.  1,  p.  167,  1941  (Japan). — 
Okada  and  Ikeda,  Biogeographica,  Trans.  Biogeogr.  Soc.  Japan,  vol.  3,  No. 
2,  p.  203,  fig.  29,  1939  (Riu  Kiu  Islands). 

Prochilus  polylepis  Bleeker,  Versl.  Akad.  Wet.  Amsterdam,  ser.  2,  vol.  11,  p.  135, 
1877;  Atlas  ichthyologique  .  .  .  ,  vol.  9,  pi.  401,  fig.  6,  1878. 

Amphiprion  polymnus  (non  Linnaeus)  Montalban,  Bur.  Sci.  Manila  Monogr. 
24,  p.  10,  pi.  1,  fig.  1,  1928  (Philippines). 

I  have  studied  72  specimens  from  the  Philippines,  Okinawa,  and 
Japan  referable  to  this  species.  In  alcohol  two  color  phases  are  evi- 
dent, the  usual  one  (53  specimens,  ranging  from  37  to  100  mm.  in 
standard  length)  has  a  plain  blackish  body  behind  head,  whereas  15, 
ranging  from  39  to  60  mm.,  have  3  lengthwise  pale  bands  on  side  of 
body,  and  4  others  are  intermediate  in  regard  to  coloration.  These 
pale  bands  at  best  are  not  very  distinct.  This  species  is  character- 
ized by  having  about  twice  as  many  of  the  specimens  with  IX  dorsal 
spines  as  with  X;  this  latter  figure  is  the  usual  number  of  dorsal 
spines  in  the  other  species  of  Amphiprion. 

Of  78  specimens  in  54  lots  studied,  76  were  from  the  Philippine 
Islands  and  one  each  from  Japan  and  Okinawa.  The  specimen  from 
Okinawa  (the  type  locality  oi frenatus),  USNM  71702,  Naha,  Okinawa, 
Albatross,  69  mm.  in  standard  length,  is  herewith  designated  as  neo- 
type  of  Amphiprion  frenatus  Brevoort  since  the  type  is  not  in  existence. 


INDO-PACIFIC    FISH    GENUS    AMPHIPRION — SCHULTZ  201 

Amphiprion  melanopus  Bleeker 

Plate  10,  Fiqubb  E 

Amphiprion  melanopus  Bleeker,  Nat.  Tijdschr.  Nederl.  Indie,  vol,  3,  p.  561,  1852. 
Prochilus  melanopus  Bleeker,  Atlas  ichthyologique  .  .  .  ,  vol.  9,  pi.  401,  fig.  7, 

1878. 
Prochilus  macrostomus  Bleeker,  Atlas  ichthyologique  .  .  .  ,  vol.  9,  pi.  401,  fig.  5, 

1878. 
Amphiprion  ephippium  (non  Bloch)   Giinther,  Journ.  Mus.  Godeflfroy,  vol.  15, 

pt.  7,  pi.  122,  fig.  D  (var.  melanopus  on  p.  225),  1881. 
Amphiprion  mccullochi  Whitley,  Mem.  Queensland  Mus.,  vol.  9,  pt.  3,  p.  213, 

1929  (Lord  Howe  Island). 
Amphiprion  macrostoma   (non  Bleeker)    Chevy,   Travaux  Inst.  Oceanogr.  Indo- 

Chine,  Mem.  4,  pt.  1,  Poissons,  p.  102,  pi.  40,  1932  (Annam). 

This  species  is  characterized  by  the  single  white  bar  on  the  head, 
plain  black  body,  pale  soft  dorsal,  caudal,  and  pectoral,  and  black 
pel  vies  and  anal  fins. 

Of  18  specimens  in  7  lots  studied,  14  were  from  the  Marshall 
Islands,  3  from  Apia,  Samoa,  and  1  from  Paumotu  Island. 


a.  t.  SOVEIMHEHT  PKINTIH*  OrFICEi  1*13 


PROCEEDINGS  OF  THE  UNITED  STATES  NATIONAL  MUSEUM 


issued  i^§y\A, m'J^I  h  the 


SMITHSONIAN  INSTITUTION 

U.  S.  NATIONAL  MUSEUM 
Vol.103  Washington:  1954  No.  3324 

MARINE  POLYCHAETE  WORMS  FROM  POINT  BARROW, 

ALASKA,  WITH  ADDITIONAL  RECORDS  FROM  THE 

NORTH  ATLANTIC  AND  NORTH  PACIFIC 


By  Marian  H.  Pettibone 


This  report  of  the  Arctic  polychaetes  found  in  the  region  of  Point 
Barrow,  Alaska,  is  based  on  material  collected  during  1948,  1949,  and 
1950  by  G.  E.  MacGinitie,  of  the  Arctic  Research  Laboratory.  Speci- 
mens were  obtained  from  Eluitkak  Pass,  Elson  Lagoon,  near  Point 
Barrow,  were  washed  ashore  at  the  Point  Barrow  base,  and  were 
dredged  within  16  miles  offshore  at  Point  Barrow  base  in  depths  of 
1.7  to  123.5  fathoms  on  bottoms  of  mud,  stones,  gravel,  rocks,  in 
masses  of  worm  tubes,  and  various  combinations  of  these.  Addi- 
tional specimens  were  collected  from  fish  traps,  from  screen  traps 
lowered  through  holes  made  in  the  ice,  and  from  plankton  hauls, 
some  of  which  were  made  through  holes  in  the  ice.  The  latter  collec- 
tions are  of  particular  interest  in  that  they  include  specimens  showing 
sexual  epitokous  stages  of  some  of  the  syllids.  Considerable  care 
was  taken  in  going  over  the  miscellaneous  material  and  separating 
polychaetes,  as  evidenced  by  the  large  collection  and  the  presence 
in  it  of  many  small  specimens  of  the  young  of  larger  species  as  well 
as  small  species  which  are  often  overlooked.  There  are  a  great  num- 
ber of  small  syllids,  phyllodocids,  hesionids,  and  terebeUids.  Great 
care  also  was  taken  in  the  preservation  of  the  specimens.  Some 
color  notes  were  included  and  some  color  photographs  were  taken,  and 
these  added  considerably  to  the  value  of  the  collection. 

The  material  was  worked  on  by  the  writer  at  the  U.  S.  National 
Museum,  where  the  collection  is  deposited.  The  facilities  of  the  Mu- 
seum, including  laboratory  accommodations,  the  library,  and  the  vast 
261112—54 1  203 


204  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  103 

polychaete  collections,  which  include  much  type  material,  were  placed 
at  the  writer's  disposal.  Some  previously  unworked  material  was 
examined  for  comparative  purposes,  and  the  results  of  some  of  these 
related  studies  are  included  here.  This  material  is  chiefly  from  collec- 
tions made  by  the  following:  R.  A.  Bartlett  in  the  Canadian  Arctic, 
Greenland,  and  Labrador  from  1927  to  1942;  W.  H.  Dall  in  Arctic 
Alaska,  the  Bering  Sea,  and  southeastern  Alaska  from  1871  to  1880; 
the  Blue  Dolphin  expeditions  to  Labrador  and  Newfoundland  from 
1949  to  1951  under  the  command  of  D.  C.  Nutt;  the  U.  S.  Fish  Com- 
mission in  dredgings  off  the  east  coast  of  North  America,  from  which 
many  specimens  had  been  identified  and  recorded  by  A.  E.  Verrill; 
and  by  the  writer  in  the  Straits  of  Juan  de  Fuca,  in  Washington  and 
Puget  Sounds,  Washington,  chiefly  during  the  summers  from  1936  to 
1940,  and  from  the  region  of  Woods  Hole,  Massachusetts,  in  the 
summers  of  1950  and  1951.  For  the  last  three  collections  mentioned, 
only  summaries  of  the  data  are  given  for  species  common  to  the 
Point  Barrow  region.  More  complete  data  are  to  be  published  sepa- 
rately and  the  summaries  are  included  here  only  to  make  the  dis- 
tributional data  of  the  Point  Barrow  species  more  complete. 

The  number  of  polychaetes  previously  recorded  from  Arctic  Alaska 
is  smaU  indeed.  The  collection  obtained  by  the  International  Polar 
Expedition  to  Point  Barrow  from  1881  to  1883  included  only  17  poly- 
chaetes (Murdoch,  1885,  p.  152).  This  collection,  which  was  depos- 
ited in  the  U.  S.  National  Museum,  was  examined  by  the  writer  and 
is  referred  to  in  this  report.  A  few  additional  records  in  scattered 
papers  have  added  to  the  list  of  polychaetes  of  Arctic  Alaska. 

For  each  of  the  species,  a  rather  full  but  by  no  means  complete 
synonymy  is  given,  chiefly  bringing  together  references  to  the  original 
and  additional  descriptions  and  scattered  distributional  records.  To 
facilitate  identification,  keys  to  the  families,  genera,  and  species  are 
given,  as  well  as  synopses  of  the  families  and  brief  descriptions  of  the 
species  with  size  ranges  and  color  notes.  An  explanatory  key  to  the 
lettering  of  the  diagnostic  features  in  the  figures  is  given  on  page  2 10. 

This  study  was  aided  by  a  contract  between  the  Office  of  Naval 
Research,  Department  of  the  Navy,  and  Johns  Hopkins  University 
(Project  No.  NR  162  911,  Contract  and  Task  Order  No.  N6onr 
243-16). 

The  writer  acknowledges  her  appreciation  to  Prof,  and  Mrs.  G.  E. 
MacGinitie  for  their  cooperation  and  help  on  this  project;  to  the  au- 
thorities of  the  U.  S.  National  Museum  for  allowing  her  to  make  use 
of  the  facilities  of  the  Institution,  especially  to  Dr.  Waldo  L.  Schmitt 
and  Dr.  Fenner  A.  Chace,  Jr.,  for  their  valuable  aid,  suggestions,  and 
patience;  and  to  Mr.  and  Mrs.  Cyiil  Berkeley,  of  Nanaimo,  British 
Columbia,  for  their  loan  of  specimens  and  their  valuable  suggestions. 


MARINE    POLYCHAETE    WORMS — PETTIBONE  205 

The  distribution  of  the  Point  Barrow  polychaetes  is  summarized 
in  table  1. 

The  collections  include  3,270  specimens  representing  88  species  and 
26  families  of  Polychaeta.  Some  of  the  species  were  exceedingly 
common;  others  were  represented  only  by  single  or  few  specimens. 
The  Polynoidae,  Syllidae,  and  Terebellidae  are  the  most  abundant 
both  as  to  numbers  of  species  (11  each,  or  37  percent)  and  specimens 
(66  percent).  The  Phyllodocidae  and  Sabellidae  have  7  species  each. 
Eleven  families  are  represented  by  a  single  species  each. 

During  1948,  because  of  the  ice  floes  all  summer,  there  was  no 
heavy  surf  and  practically  nothing  was  washed  up  on  the  beach. 
During  1949,  specimens  were  washed  ashore  on  17  different  days. 
In  1950,  specimens  were  collected  from  the  beach  on  only  one  day. 
Altogether,  287  specimens  of  polychaetes  were  washed  ashore,  repre- 
senting 20  species  and  11  families.  Two  species,  Eunoe  clarki  (2 
specimens)  and  Travisia  carnea  (12  specimens),  were  obtained  in  this 
manner  only.  Four  species  were  dredged  occasionally  but  were 
washed  ashore  in  much  larger  numbers,  namely:  Antinoe  sarsi  (58 
specimens),  Alelaenis  loveni  (65  specimens),  Arenicola  glacialis  (54 
specimens),  and  Brada  villosa  (24  specimens).  Additional  species 
were  commonly  obtained  in  the  dredge  but  were  washed  ashore  in 
small  numbers. 

On  three  occasions  in  1950  when  80-foot  vertical  plankton  hauls 
were  taken  through  holes  in  the  ice,  the  sexual  stages  of  three  of  the 
syllids  were  found — Avtolytits  fallax,  Syllis  cornuta,  and  Syllis  fasciata. 

A  station  12.1  miles  from  shore,  123.5  fathoms,  August  17,  1949, 
produced  literally  bushels  of  worm  tubes.  The  tubes  of  the  terebellid 
Pista  maculata  made  up  the  greater  part  of  the  mass. 

Of  the  88  Point  Barrow  species,  75  (85.2%)  are  common  to  the 
Arctic,  Atlantic,  and  Pacific;  5  (5.7%)  are  common  only  to  the  Arctic 
and  Pacific  {Gattyana  ciliata,  Eusyllis  magnijica,  Pionosyllis  compacta, 
Glycinde  wireni,  and  Idanthyrsus  armatus);  3  (3.4%)  are  common  to 
the  Arctic  and  Atlantic  {Autolytus  fallax,  Eumida  minuta,  Travisia 
carnea);  4  (4.5%)  are  confined  to  the  Arctic  (Eunoe  clarki,  Nerinides 
sp.,  Arenicola  glacialis,  Ampharete  vega);  and  1  (1.1%)  is  bipolar, 
known  only  from  the  Arctic  and  Antarctic  {Ammotrypane  hreviata). 

The  Point  Barrow  records  help  to  complete  the  circumpolarity 
records  for  58  circumpolar  or  almost  circimipolar  species,  some  of 
which  previously  had  been  known  only  from  the  Siberian  and  Canadian 
Arctic,  Greenland,  and  Spitsbergen.  The  range  has  been  extended 
for  28  species,  6  of  which  have  been  extended  by  combining  Atlantic 
and  Pacific  specific  names. 


206 


PROCEEDINGS    OF   THE    NATIONAL   MUSEUM 


■«*  -5 


d 

1 
o 

3 
& 

O 

5" 

1     1 

6  a     to- 
rt =«--  ""C3 

X 

1           X           X 

Aretic- 
boreal- 

lusi- 
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(18 
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20%) 

I 

X 

X 

K    X 

X    X     X     X 

XXX 

X 

X 

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in 

masses  of 

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gravel 

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bottom  of 

stones, 

gravel, 

pebbles, 

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(51  species) 

K 

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X     X    X    X    X 

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p§^    1 

>< 

X    X 

X    X      1 

X     X     X     X 

X 

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X 

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Number 
of 
speci- 
mens 

CS    CO                     1-H 

lis 

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Gattyana  ciliata  Moore 

SIGALIONIDAE  (30  Specimens) 

Pholo6  minuta  (Fabricius) 

3     1 
11 

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1 

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Eteone  flava  (Fabricius) 

Eteone  spetsbergensis  Malmgren 

Phyllodoce  groenlandica  Oersted 

Eumlda  minuta  (Ditlevsen) 

1 

1 

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11 

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MARINE    POLYCHAETE    WORMS — PETTIBONE 


207 


||]«||i;]l|        X     \        xlll        «          1          1        X          !lll        XX        X   X     \ 

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«|H|Kt<!|||K|              II              ',     X    X     X              1              1              1              1            X       [     X       ',              11              III 

]    X       ',       \       \       \       \       ',       \       ',    X              \    X              llll              1           X              1              1              IIIH              II              ll« 

X       \       \    X     X       ',    X       \     X       ',    X              \    X              ',       \     X    X              1              1            X              1              llll              IH           X       [     X 

XXXXXXXXX\X            XX              |««|           X            X           K            1«)           X     X    X       \           XX           XXX 

X    X    X    X    X       \    X      \    X    X       \             11           X    X    X       \             1           X           X           X           X      \       \      \           XX           X    X      \ 

1h|1x|1I1||          II          \   X   X   X          1          1        X        X          llll          11          III 

lx|lllllx||           11         X      \    X      \           1           1           1           1           llll           II         X    X      \ 

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208 


PROCEEDINGS    OF    THE    NATIONAL   MUSEUM 


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1 

3 
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3 
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1       X     1         1         1         !    !    !    I    I         1         I         1    !         1     !    !    1 

Cosmo- 
politan 

(21 

species, 

24%) 

X         11       X         1        ]  X  X    \    ',         !       X         ;;        1111 

Arctic- 
boreal- 

lusi- 
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(18 
species, 

20%) 

1        11        1        1       xllll       X        ;        ;i        l«ll 

Arctic- 
boreal 

(30 

species, 

34%) 

1            [    X            1            1            \      ',      \    X    X            1            ]          XX            1111 

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Arctic 

(18 

species, 
20%) 

1         11         1       X         1111!        1         1        11       X    \  X  X 

a 

3 
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(35 

species) 

X         11         1         1         ',    ',  X    ',  X        ;         ;       XX         11«1 

Dredged 

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stones, 

gravel 

(74 
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K           X       ',            X              1           X       ',     X    X       \            X              1           XX              ',    X    X       \ 

Dredged  on 

bottom  of 

stones, 

gravel, 

pebbles, 

with  mud 

(51  species) 

X             ;l             1           X           X    X       ',      \       \             ;           X           XX             ',    X    X      \ 

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(31 
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mens 

t^           C^C<            O           t^           O»-H00»Oi-^            CO           "3           ^^           *^^S5^ 

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c^        fjM        <N        «        coMcococc        CO        CO        coco        coeococo 

1 

CO 

1 

1 

ScALiBREGMiDAE  (17  Specimens) 

Scalibregma  inflatum  Rathke 

Ophelhdae  (14  specimens) 

Ammotrypane  breviata  Ehlers 

Travisia  carnea  Verrill... 

CAPrTELLiDAE  (26  Specimens) 

Capitella  capitata  (Fabricius).. 

Akenicolidae  (67  specimens) 

Arenicola  glacialis  Murdoch.. 

Maldanidae  (35  specimens) 

Praxillella  praetermissa  (Malmgren).. 

Maldane  sarsi  Malmgren 

Nicomache  lumbricahs  (Fabricius) 

Nicomache  personata  Johnson 

Petaloproctus  tenuis  (Thfiel) 

Sabellarudae  (6  specimens) 

Idanthyrsus  armatus  Kinberg.. 

Sternaspidae  (5  specimens) 

Stemaspis  scutata  (Ranzani)... 

Pectinarhdae  (72  specimens) 

Pectinaria  granulata  (Linnfi) 

Pectinaria  hyperborea  (Malmgren) 

Ampharetidae  (41  specimens) 

Ampharete  vega  (Wirfen) 

Ampharete  acutifrons  (Grube) 

Ampharete  gogsi  Malmgren 

Asabellides  sibirica  ( Wirgn) 

MARINE    POLYCHAETE    WORMS — PETTIBONE 


209 


M    M     «     X    X 


>i    X    X    X 


X     X     H     X     X 


rf    1-H    OS    00    CO    f-H    -* 


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i-i    (N    i-i    CO    W 


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cocococccocccococococc 


lO  CO  00  Ol  Oi  o 
CO  CO  CO  CO  CO  ^ 
CO  CO  CO  CO  CO  CO 


210 


PROCEEDINGS    OF   THE    NATIONAL   MUSEUM 


Explanation  of  symbols  on  figures 

Roman  numerals  indicate  body  segments. 


a,  anus 

aBr,  anal  branchia 

aC,  anal  cirrus 

aCo,  anal  cone 

aCy,  anal  cylinder 

aK,  anal  knob 

aLi,  anal  ligule 

abd,  abdominal  region 

ac,  aciculum  (heavy  spine  buried  in 
parapodial  ramus) 

acH,  acicular  genital  hook 

acL,  acicular  lobe 

ach,  achaetous  lobe 

ai,  aileron  of  jaw 

an,  antenna 

antLa,  anterior  or  presetal  lamella 

anta,  antanal  achaetous  segment 

ba,  barbule 

bl,  blade  of  seta  (terminal  or  end 
piece  of  composite  or  jointed  seta) 

br,  branchia 

brF,  branchial  filament 

brLo,  branchial  lobe 

brP,  brood  pouch 

buS,  buccal  segment 

cP,  cephalic  peak 

cCre,  cephalic  crest 

cPl,  cephalic  plate 

cR,  cephalic  ridge 

caPl,  calcareous  plate 

CO,  collarette 

cph,  cirrophore  (base  of  a  tentacular 
cirrus  or  a  dorsal  cirrus) 

cr,  crotchet 

ere,  crest 

dC,  dorsal  cirrus 

dL,  dorsal  lobe 

dN,  dorsal  notch 

dTu,  dorsal  tubercle  (lobe  correspond- 
ing to  elytrophore  on  non-elytra- 
bearing  segment) 

deR,  denticled  rim 

Dor,  dorsal 

el,  elytron  or  scale 

elph,  elytrophore  (lobe  bearing  ely- 
tron) 

ey,  eye-spot 

fTu,  facial  tubercle 

frAn,  frontal  antenna 


frH,  frontal  horn 

h  jaw 

lAn,  lateral  antenna 

IGr,  lateral  groove 

ILo,  lateral  lobe 

IN,  lateral  notch 

lOr,  lateral  sensory  organ 

liPl,  limbate  plate 

loL,  lower  lip 

mAn,  median  antenna 

mLo,  median  lobe 

maxR,  maxillary  or  distal  ring 

mo,  mouth 

nPa,  nephridial  papilla 

ne,  neuropodium  or  ventral  ramus 

neC,  neuropodial  cirrus 

neLi,  neuropodial  ligule 

neS,  neuroseta  (seta  of  neuropodium) 

neTo,  neuropodial  torus 

no,  notopodium  or  dorsal  ramus 

noLi,  notopodial  ligule 

noS,  notoseta  (seta  of  notopodium) 

noPi,  notopodial  pinnule 

noTo,  notopodial  torus 

nuEp,  nuchal  epaulette 

nuF,  nuchal  fold,  lobe  or  collar 

nuG,  nuchal  groove 

nuH,  nuchal  hook 

nuO,  nuchal  organ 

nuPa,  nuchal  papilla 

oR,  oral  or  basal  ring 

oT,  oral  tentacles 

ocT,  occipital  tubercle 

op,  operculum 

opPe,  opercular  peduncle 

p,  palp 

pM,  palmar  membrane 

pa,  papilla 

pal,  paleae 

parF,  parapodial  flange 

para,  paragnaths  or  horny  denticles 

parath,  parathoracic  segment 

poL,  postsetal  or  posterior  lobe 

poLa,  postsetal  or  posterior  lamella 

pr,  prostomium 

preL,  presetal  or  anterior  lobe 

pro,  proboscis 

pyF,  pygidial  or  anal  funnel 

pyPl,  pygidial  plate 


MARINE    POLYCHAETE    WORMS — PETTIBONE 


211 


s,  seta  uL,  upper  lip 

scH,  scaphal  hook  un,    uncinus    (very   small   and   short, 

set,  setigerous  segment  or  setiger  flattened  setal  hooks) 

setLo,  setigerous  lobe  unPi,  uncinigerous  pinnule  (flattened 

sh,  shaft  of  seta  (basal  part  of  com-  parapodial  ramus  bearing  numerous 

posite  or  jointed  seta)  uncini  or  small  setal  hooks) 

spH,  special  hook  unTo,    uncinigerous    torus    (low,    in- 

tC,  tentacular  cirrus  flated    parapodial    ramus     bearing 

tF,  tentacular  filament  numerous     uncini    or    small    setal 

tM,  tentacular  membrane  hooks) 

tS,  tentacular  segment  or  peristomium  vC,  ventral  cirrus 

(anterior  modified  segments  around  vF,  ventral  faecal  groove 

mouth)  vL,  ventral  lobe 

th,  thoracic  region  vPa,  ventral  papilla 

thM,  thoracic  membrane  vSh,  ventral  shield 

to,  tooth  Ven,  ventral 

tor,    torus    (low,    inflated    parapodial  w,  basal  crenulate  wing 

lobe) 

Order  POLYCHAETA 
Key  to  the  families  of  Polychaeta  from  Point  Barrow 

1.  Dorsal  surface  more  or  less  completely  covered  by  paired  overlapping  scales 

or  elytra  (fig.  26,  a) 5 

Dorsal  surface  not  covered  by  elytra 2 

2.  Anterior  end  more  or  less  concealed  by  feathery  tentacles  (fig.  38,  a),  chiti- 

nous  golden  setae  forming  an  operculum  (figs.  34,  n;  35,  d),  long  setae 
directed  forward  forming  a  cephahc  cage,  or  long  filamentous  outgrowths 

(fig.  36,  /) .     With  more  or  less  permanent  tubes 6 

Anterior  end  otherwise 3 

3.  Body  thickly  papillated,  grub-shaped 21 

Body  smooth  or  finely  granulated 4 

4.  Prostomium  with  conspicuous  antennae,  with  or  without  fleshy  palps  (figs. 

26-30),  or  prostomium  or  an  anterior  segment  with  pair  of  long,  prehensile, 

tentacular  palps  (fig.  32,  i) 13 

Prostomium  without  palps  and  without  conspicuous  antennae  (fig.  32,  a)  or 
with  2  pairs  minute  antennae  (figs.  30,  j;  31,  o,  e) 18 

5.  Elytra  on  all  segments  of  posterior  region  (at  least  from  segment  29  on). 

Dorsal  cirri  lacking Sigalionidae  (p.  228) 

Elytra  not  on  all  segments  of  posterior  region.     With  paired  dorsal  cirri  on 
segments  without  elytra  (fig.  26,  a) Polynoidae  (p.  213) 

6.  Anterior  end  with  branchial  plume  (fig.  38,  a) 7 

Anterior  end  without  branchial  plume 8 

7.  Branchial    filaments    without    operculum.     Without    thoracic    membrane. 

Tube  gelatinous,  membranous,  or  horny,  covered  or  not  with  mud,  sand, 

gravel,  and  debris Sabellidae  (p.  332) 

Usually  1  or  2  opercula  on  modified  branchial  filaments  (fig.  39,  u) .     Usually 
with  thoracic  membrane  (fig.  39,  u).   Tube  calcareous.. Serpulidae  (p.  342) 

8.  Anterior  end  with  chitinous,  golden  setae  forming  an  operculum  (figs.  34, 

n;  35,  d).     In  free,  conical,  sandy  tubes  or  concreted  sandy  tubes  attached 

to  rocks,  shells,  etc 12 

Anterior  end  without  opercular  setae.     Tube  otherwise 9 


212  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  103 

9.  Setae  of  anterior  segments  directed  anteriorly,   forming  a  cephalic   cage. 
Body  covered  with  papillae  and  often  with  adherent  sand  grains;  may  be 

encased  in  thick  mucoid  sheath Flabelligeridae  (p.  288) 

Anterior  end  without  cephalic  cage  but  with  long,  filamentous  outgrowths 
(may  have  2  groups  of  paleal  setae  directed  forward;  fig.  36,  a,  e,  /).  Body 
without  papillae 10 

10.  Anterior  end  with  long,  filamentous  outgrowths  and  with  similar  outgrowths 

continued  along  sides  of  body  on  few  to  many  segments  (fig.,  33,  a).    Seg- 
ments similar,  body  not  divided  into  2  distinct  regions. 

Cirratulidae  (p.  285) 

Long,  filamentous  outgrowths  concentrated  on  anterior  region  only  (figs.  36, 

37).     Body  with  2  distinct  regions 11 

11.  With  numerous  filiform  oral  tentacles  retractile  in  mouth  (fig.  36,  e).     Ante- 

rior region  with  or  without  2  groups  of  paleal  setae  directed  forward  (fig. 

36,  a,  b) Ampharetidae  (p.  314) 

With  numerous,  grooved,  oral  tentacles  not  retractile  in  mouth  (fig.  36,  /,  g, 
i) .     Anterior  region  without  paleal  setae Terebellidae  (p.  318) 

12.  Opercular  setae  forming  1-3  concentric  rows   (fig.  34,  n-q).     Tube  of  con- 

creted sand  or  fine  gravel,  attached Sabellariidae  (p.  307) 

Opercular  setae  forming  2  bundles  in  horizontal  rows,  directed  obliquely 
anteriorly  (fig.  35,  c,  d).  Tube  free,  conical,  rigid,  open  at  both  ends, 
formed  of  single  layer  of  sand  grains Pectinariidae  (p.  310) 

13.  Prostomium  with  2-5  conspicuous  antennae.     With  1-8  pairs  tentacular  cirri 

(figs.  27-29) 14 

Prostomium  without  conspicuous  antennae.  Without  tentacular  cirri.  With 
pair  of  tentaculiform,  longitudinally-grooved  palps  (deciduous,  may  be 
missing;  figs.  32,  i;  33,  d) 17 

14.  Dorsal  cirri  flattened,  foliaceous,  sometimes  overlapping,  or  more  or  less 

globular    (fig.    27).     Prostomium    with    2    eyes,    4-5    antennae,    without 

palps Phyllodocidae  (p.  231) 

Dorsal  cirri  otherwise.  Prostomium  ^vith  4-6  eyes,  2-3  antennae,  2  palps 
(may  be  reduced  and  fused  in  Syllidae) 15 

15.  With  parapodial  ligules,  rounded,  conical  or  straplike  lobes  (fig.  30,  h,  c). 

Dorsal  cirri  short,  cirriform Nereidae  (p.  263) 

Without  parapodial  ligules.  Dorsal  cirri  filiform,  may  be  more  or  less  articu- 
lated (or  small  and  subulate  in  some  Syllidae) 16 

16.  Palps  biarticulate  (fig.  28,  a) Hesionidae  (p.  239) 

Palps  not  biarticulate  (maj^  be  reduced  and  fused;  figs.  28,  c;  29,  a). 

Syllidae  (p.  240) 

17.  Parapodia  with  postsetal  lamellae  more  or  less  developed  (fig.  32,  g).     Branch- 

iae  usuall.y  paired,   dorsal,   ligulate   or   cirriform    (rarely  pinnate,  rarely 

lacking) Spionidae  (p.  280) 

Parapodia  with  lobes  scarcely  projecting,  without  postsetal  lamellae.  Branch- 
iae simple,  filamentous,  contractile,  inserted  above  notopodia  on  few  to 
many  segments  (fig.  33,  d) Cirratulidae  (p.  285) 

18.  Prostomium  with  2  pairs  minute  antennae  (figs.  30,  j;  31,  o) 19 

Prostomium  without  antennae 22 

19.  Prostomium  subquadrate,  with  small  antennae  on  anterior  part  (fig.  30,  j). 

With  anterior  and  posterior  parapodial  lamellae  more  or  less  developed 

(fig.  30,  k-n) Nephtyidae  (p.  266) 

Prostomium  conical,  pointed,  annulated,  with  tiny  terminal  antennae  (fig. 
31,  o,  e).     Without  parapodial  lamellae 20 


MARINE    POLYCHAETE    WORMS — PETTIBONE  213 

20.  Segments  bi-  or  triannulate.     Parapodia  of  single  form  throughout  length  of 

body,  either  only  uniramous  or  only  biramous Glyceridae  (p.  272) 

Segments  uniannulate.  Body  divided  into  2  regions — anterior  uniramous, 
posterior  biramous  (fig.  31,/,  g) Goniadidae  (p.  274) 

21.  Posterior  part  of  body  with  2  horny  ventral  plates  with  radiating  bundles  of 

setae  (fig.  35,  6) Sternaspidae  (p.  309) 

Posterior  part  of  body  without  horny  plates Flabelligeridae  (p.  288) 

22.  Some  of  segments  considerably  elongated,  much  longer  than  wide.     Anal 

segment  with  limbate  plate,  spatulate,  or  funnellike   (fig.  34,  d,  h,  rn). 
Prostomium  hoodlike  (fig.  34,  i,  I)  or  with  limbate  plate  (fig.  34,  b-c,  g). 

Maldanidae  (p.  302) 
Segments  not  much  longer  than  wide 23 

23.  Proboscis  provided  with  dark,  chitinous  jaw  pieces  (fig.  31,  i).     Body  smooth, 

elongate,  cylindrical,  resembling  an  earthworm.     Parapodia  weakly  devel- 
oped simple  lobes Lumbrineridae  (p.  275) 

Proboscis  without  chitinous  jaws.     Form  variable 24 

24.  Body  divided  into  2  weakly  to  sharply  separable  regions,  short  anterior  region, 

more  or  less  flattened,  with  cushionlike  neuropodia  with  several  rows  of 
setae,  and  long  cylindrical  posterior  region  with  parapodial  lobes  projecting 

dorsally  (fig.  32,  c) Orbiniidae  (p.  278) 

Parapodial  lobes  of  median  and  posterior  regions  not  projecting  dorsally..  25 

25.  Prostomium  conical 26 

Prostomium  otherwise 27 

26.  Body  short,  grub-shaped  or  lancet-shaped.     Parapodia  with  simple  capillary 

setae  only Opheliidae  (p.  295) 

Body  elongate,  slender,  fragile.  Parapodia  with  capillary  setae  and,  in  long 
posterior  region,  with  rows  of  hooded  hooks  on  somewhat  inflated  tori 
(fig.  33,  t) Capitellidae  (p.  298) 

27.  Arborescent  branchiae  limited  to  anterior  few  segments.     With  capillary  and 

forked  parapodial  setae Scalibregmidae  (p.  293) 

Arborescent  branchiae  on  long  middle  region  of  body.  With  capillary  setae 
and  sigmoid  hooks  embedded  in  elongated  tori Arenicolidae  (p.  300) 

Family  Polynoidae 

Prostomium  bilobed,  with  two  pairs  of  eyes  in  trapezoidal  arrange- 
ment, three  dorsal  antennae,  a  pair  of  subulate  ventral  palps.  First 
or  tentacular  segment  with  setae  reduced  in  number  (0-3),  with  two 
pairs  of  tentacular  cirri  (fig.  26,  a).  Parapodia  biramous,  with  setae 
simple  (not  compound),  paired  dorsal  scales  or  elytra  on  certain  seg- 
ments and  paired  dorsal  cirri  on  non-elytra-bearing  segments,  paired 
subulate  ventral  cirri  (fig.  26,  a,  d).  Pygidium  or  anal  segment  with 
pair  of  anal  cirri.  Paired  segmental  or  nephridial  papillae  near 
ventral  bases  of  parapodia.  Muscular  proboscis  eversible,  with  circle 
of  soft  marginal  papillae  and  two  pairs  of  interlocking,  chitinous  jaws. 

Represented  by  7  genera  and  11  species.  All  the  genera  con- 
sidered here  have  15  pairs  of  elytra  on  segments  2,  4,  5,  7,  9,  11,  13, 
15,  17,  19,  21,  23,  26,  29,  32;  lateral  antennae  inserted  ventral  to 
median  antenna  on  prostomium  (fig.  26,  a). 


214  PROCEEDINGS    OF   THE   NATIONAL   MUSEUM  vol,,  los 

Key  to  the  genera  of  Polynoidae  from  Point  Barrow 

1.  Notosetae  stouter  than  or  at  least  as  stout  as  neurosetae,  with  tips  blunt  to 

pointed,  not  capillary 2 

Notosetae  not  as  stout  as  neurosetae 5 

2.  Notosetae  large,  smooth  or  with  only  faint  transverse  striations,  few  in  number 

(5-9) Melaenis  (p.  214) 

Notosetae  with  transverse  spinous  rows,  more  numerous  (more  than  10) 3 

3.  Neurosetae  long,  slender,  at  least  some  end  in  capillary  tips.Antinoe  (p.  215) 
Neurosetae  stouter,  with  tips  straight  or  slightly  hooked,  not  capillary 4 

4.  All  neurosetae  with  entire  bare  tips Eunoe  (p.  216) 

Some  of  neurosetae  bidentate,  with  secondary  tooth  well  developed  or  rudi- 
mentary   (fig.   26,   e) Harmothoe    (p.   220) 

5.  Some  notosetae  with  capillary  tips.     Segments  less  than  40  (35-38),  thus, 

posterior  end  of  body  covered  with  elytra 6 

None  of  notosetae  with  capillary  tips.  Segments  more  than  40  (45-65),  thus, 
a  rather  long  posterior  end  without  elytra.  Elytra  smooth,  without 
tubercles Enipo  (p.  225) 

6.  Neurosetae  of  2  kinds,  upper  few  end  in  slender  tips,  rest  end  in  bifid  tips. 

Elytra   smooth   except   for  few   microtubercles   on   anterior  curved  part. 

Arcteobia  (p.  225) 

Neurosetae  all  similar,   with  entire   tips.     Elytra  with   microtubercles  and 

sometimes  also  with  macrotubercles Gattyana  (p.  622) 

Genus  Melaenis  Malmgren,  1865 
Melaenis  loveni  Malmgren,  1865 

Melaenis  lovini  Malmgren,  1865,  p.  78,  pi.  10,  fig.  10.— Th^el,  1879,  p.  22.— 
Wir^n,  1883,  p.  391,  pi.  28,  fig.  4;  pi.  29,  fig.  3.— Murdoch,  1885,  p.  152.— 
Moore,  1908,  p.  337.— Fauvel,  1914,  p.  49.— Augener,  1928,  p.  695.— 
Ditlevsen,  1937,  p.  16.— Gorbunov,  1946,  p.  38.— Wesenberg-Lund,  1950a, 
p.  9;  1950b,  p.  29. 

Melaenis  lov6ni  var,  gigantea  Wir4n,  1883,  p.  391,  pi.  28,  fig.  3;  pi.  29,  fig.  4. — 
Murdoch,  1885,  p.  153. 

Description. — ^Ijength  25-76  mm.,  width  including  setae  9-22  mm- 
(up  to  110  mm.  long — Ditlevsen,  1937).  Segments  39-41.  Body 
elongated,  thick,  tapering  both  anteriorly  and  posteriorly.  Pro- 
stomium  without  cephalic  peaks,  antennae  smooth.  Dorsal  cirri 
with  fine,  scattered,  clavate  papillae.  Elytra  smooth,  soft,  trans- 
lucent to  opaque,  not  covering  middorsum  and  posterior  few  segments. 
Notosetae  much  stouter  than  neurosetae,  few  in  number  (5-9), 
light  to  dark  amber-colored,  smooth  or  with  only  very  faint  transverse 
striations.  Neurosetae  of  two  kinds:  Numerous,  long,  slender, 
finely  denticled,  with  capillary  tips;  few,  shorter,  with  subequal, 
diverging,  forked,  blunt  tips  (one  part  of  fork  may  be  longer  than  the 
other).  Color:  In  life  and  in  alcohol:  Wide  greenish  brown  to 
bluish  gray  transverse  stripes  middorsally,  green  and  white  striped  on 
dorsal  tubercles,  cirrophores  and  elytrophores;  without  color  ventrally; 
elytra  light  to  dark  greenish  in  a  wide,  circular  band,  a  circular  area 


MARINE    POLYCHAETE    WORMS — PETTIBONE  215 

and  lateral  border  without  color;  one  specimen  (collected  October  4, 
1949)  was  noted  as  peach-colored  from  segment  10  posteriorly,  the 
color  showing  mainly  on  the  ventral  side  and  extending  into  the 
parapodia  (probably  developing  eggs). 

New  records. — Arctic  Alaska:  Point  Barrow  base,  washed  ashore, 
1949  (July  30;  August  21;  September  22,  28,  30;  October  4,  5,  16, 
17;  65  specimens);  off  Point  Barrow  base,  along  shore,  1.7  fms.,  on 
bottom  of  gravel  with  mud  (2  specimens).  East  Coast  North 
America:  Off  Labrador,  45  fms.,  Blue  Dolphin  Expedition,  1949. 

Distribution.— Widely  distributed  m  the  Arctic:  Siberian  and 
Alaskan  Ai'ctic,  Davis  Strait,  Greenland,  Spitsbergen,  Franz  Josef 
Land,  Kara  Sea.  Also  Bering  Sea;  off  Labrador.  In  1.7-111 
fathoms. 

Genus  Antinoe  Kinberg,  1855  (sensu  Malmgren,  1865) 
Antinoe  sarsi  Malmgren,  1865 

Antinoe  sarsi  Kinberg,  1862  (MS.),  p.  468  (fide  Malmgren,  1865).— Malmgren, 

1865,  p.  75,  pi.  9,  fig.  6  (part)  .—Mcintosh,  1900,  p.  365,  figs.— Chamberlin, 

1920,  p.  8. — Hartman,  1944a,  p.  334. 
Polynoe  sarsi  Theel,  1879,  p.  16,  pi.  1,  fig.  8.— Wir6n,  1883,  p.  390.— Murdoch, 

1885,  p.  152. 
Harmothoe  sarsi  Eliason,   1920,  p.   20.— Annenkova,   1931,  p.   203.— Friedrich, 

1939,  p.  122.— Thorsou,  1946,  p.  48.— Wesenberg-Lund,  1950a,  p.  7;  1950b, 

p.  23;  1951,  p.  17. 
Harmothoe  (Antinoella)  sarsi  Augener,  1928,  p.  687. 
Antinoella  sarsi  Annenkova,  1937,  p.  153;  1938,  p.  137.— Gorbunov,  1946,  p.  38.— 

Zatsepin,  1948,  p.  107,  pi.  28,  fig.  8. 
Description.— Length.  21-68  mm.,  width  mcluding  setae  11-27  mm. 
Segments  37  or  38.  Fragments  easily,  posterior  segments  frequently 
regenerating.  Body  elongate,  flattened,  tapering  shghtly  anteriorly 
and  posteriorly.  Prostomium  with  cephalic  peaks  distinct  or  poorly 
developed;  anterior  pan-  of  eyes  larger  than  posterior,  slightly  anterior 
to  greatest  width  of  prostomium.  Antennae  with  short,  clavate 
papillae.  Dorsal  cirri  extending  beyond  setae,  long,  slender,  tapering 
gradually,  with  short,  clavate  papillae.  May  be  a  single  anal  ckrus, 
extremely  long,  thick  at  base,  tapering  gradually;  may  be  one  long 
and  one  short  anal  cirrus.  Elytra  large,  thin,  soft,  smooth,  with 
scattered  microtubercles  and  short,  delicate,  clavate  papillae.  Lose 
elytra  readily;  often  show  regenerating  elytra,  thus  appearing  variable 
in  size.  Notopodia  and  neuropodia  extending  into  conspicuous, 
digitiform,  acicular  lobes.  Setae  yellowish  or  golden.  Notosetae 
much  thicker  than  neurosetae,  large,  clear  basally;  more  distally, 
finely  spinous,  pointed.  Neurosetae  of  two  kinds,  with  capillary  tips 
and  with  slender,  relatively  obtuse,  shghtly  curved  tips.  Color: 
In  life  and  preserved:  Grayish  gi-een  or  light  brown  middorsally, 


216  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  i03 

including  elytrophores  and  dorsal  tubercles;  ventral  surface  without 
color  or  dusky  midventrally;  elytra  gray,  translucent  to  opaque, 
with  a  darker  brownish-gray  medial  crescent-shaped  area,  with  a 
colorless  circular  area  medial  to  and  a  darker  area  posterior  to  the 
place  of  attachment. 

Remarks. — A.  sarsi  differs  from  A.  badia  (Theel),  with  which  it  has 
sometimes  been  confused,  by  having  the  neurosetae  of  two  lands, 
with  capillary  tips  and  with  slender,  relatively  obtuse  tips.  In  A. 
badia,  all  the  neurosetae  have  capillary  tips. 

Parasites. — One  of  the  63  specimens  had  the  parasitic  copepod 
Herpyllobius  arcticus  Steenstrup  and  Liitken  attached  to  the  prosto- 
mium  (identified  by  P.  L.  Illg). 

New  records. — Arctic  Alaska:  Point  Barrow  base,  washed  ashore 
1949  (August  21,  24;  September  1,  12,  20,  22,  24,  26;  October  5,  16; 
58  specimens) ;  off  Point  Barrow  base,  up  to  5  miles  from  shore,  5  to 
30.7  fms.,  on  bottoms  of  mud  or  stones,  and  in  screen  trap  through 
hole  in  ice,  6  fms.,  April  11,  1949  (3  stations,  5  specimens);  near 
Point  Belcher,  Icy  Cape,  off  Point  Barrow,  Dall.  Bering  Sea:  62°15' 
N.,  167°48'  W.,  20  fms.,  G.  M.  Stoney,  1884.  East  Coast  North 
America:  Off  Labrador,  6-30  fms.,  Blue  Dolphin  Expeditions, 
1950,  1951. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian  and 
Alaskan  Arctic,  Greenland,  Jan  Mayen,  Spitsbergen,  Norway,  Franz 
Josef  Land,  Barents  Sea,  Novaya  Zemlya,  Kara  Sea.  Also  Iceland, 
Faroes  to  Great  Britain,  the  Baltic;  Labrador  to  Maine;  Bering  Sea; 
north  Japan  Sea.     In  3-1,215  fathoms. 

Genus  Eunoe  Malmgren,  1865 
Key  to  the  species  of  Eunoe  from  Point  Barrow 

1.  Elytra  with  scattered,  bluntly  conical  microtubercles  only,  lacking  fringe  of 

papillae.  Eyes  small,  anterior  pair  anterolateral  on  prostomium  (scarcely 
visible  dorsally) E.  clarki 

Elytra  with  macrotubercles  in  addition  to  microtubercles,  with  lateral  fringe 
of  papillae.     Eyes  larger,  anterior  pair  anterodorsal  (visible  dorsally) 2 

2.  Body  broad,  oval,  flattened.     Extra  rounded  lobes  on  inner  sides  of  elytro- 

phores and  dorsal  tubercles  (lobes  corresponding  to  elytrophores  on  non- 
elytra-bearing  segments,  fig.  26,  c).  Nuchal  fold  posterior  to  prostomium 
not  prominent.  Setae  yellow.  Elytra  with  microtubercles  rather  low, 
flattened,  semiglobose,  some  bifid;  macrotubercles  confined  essentially  to 
single  row  near  external  border,  nodular,  with  roughened  tips.     Antennae 

with  short  papillae E.  nodosa 

Body  more  elongate,  not  so  flattened.  Without  extra  rounded  lobes  on  inner 
sides  of  elytrophores  and  dorsal  tubercles.  Nuchal  fold  prominent.  Setae 
dark  amber-colored.  Elytra  with  microtubercles  one-  to  many-pronged; 
macrotubercles  variable  in  number,  size,  and  position,  branched.  Antennae 
with  longer  papillae E.  oerstedi 


MARINE    POLYCHAETE    WORMS — PETTIBONE  217 

Eunoe  clarki  Pettibone,  1951 

Eunoe  clarki  Pettibone,  1951,  p.  44,  fig.  1. 

Description. — ^Length  36-38  mm.,  width  including  setae  12  mm. 
Segments  40  or  41.  Body  linear-oblong,  tapered  slightly  anteriorly 
and  posteriorly.  Prostomium  without  cephalic  peaks ;  anterior  pair  of 
eyes  anterolateral.  Antennae  and  dorsal  cirri  with  short,  clavate 
papillae.  Elytra  nearly  cover  dorsum,  with  conical  micro  tubercles, 
without  fringe  of  papillae.  Color:  In  alcohol:  Middorsum  trans- 
versely banded  grayish  green;  elytra  greenish  gray,  with  a  darker 
mottled  pattern  on  most  of  the  exposed  parts,  with  a  darker  spot 
medial  to  a  lighter  area  over  the  elytophore;  dorsal  cirri  with  pig- 
mented bands  basally  and  subterminally. 

New  record. — Arctic  Alaska:  Point  Barrow  base,  washed  ashore, 
October  17,  1949  (2  specimens). 

Distribution. — Arctic  Alaska. 

Eunoe  nodosa  (Sars,  1860) 

Figure  26,  c 

Polynoe  nodosa  Sars,  1860,  p.  58. 

Eunoe  nodosa  Malmgren,  1865,  p.  64,  pi.  8,  fig.  4. — Fauvel,  1923,  p.  51,  fig.  18, 

a-e  (part).— Annenkova,  1937,  p.  149;  1938,  p.  134.— Treadwell,  1937,  p. 

27.— Friedrich,  1939,  p.  122.— Berkeley  and  Berkeley,  1942,  p.  187;  1943, 

p.  129.— Hartman,  1944a,  pp.  334,  337.— Gorbunov,  1946,  p.  38.— Zatsepin, 

1948,  p.  107,  pi.  28,  fig.  7. 
Eunoa  nodosa  Malmgren,  1867,  p.  6. — Webster  and  Benedict,  1884,  p.  700;  1887, 

p.  708.— Mcintosh,  1900,  p.  291,  figs. 
Polynoe  scabra  Th^el,  1879,  p.  7.— Wir6n,  1883,  p.  387,  pi.  28,  fig.  2;  pi.  29,  fig. 

1  (part;  not  pi.  28,  fig.  1;  not  Aphrodita  scabra  Fabricius,  1780). 
Polynoe  islandica  Hansen,  1882,  pp.  17,  24,  pi.  1,  figs.  15-21.— Murdoch,  1884, 

p.  152. 
Polynoe  arctica  Hansen,  1882,  pp.  21,  27,  pi.  3,  figs.  1-5. 
Polynoe  spinulosa  Hansen,  1882,  p.  28,  pi.  1,  figs.  6-10. 
Polynoe  foraminifera  Hansen,  1882,  pp.  21,  23,  29,  pi.  1,  figs.  11-14. 
Harmothoe  nodosa  Moore,  1902,  p.  271. — Ditlevsen,  1917,  p.  6,  pi.  3,  fig.  10  (part; 

not  pi.  2,  fig.  1).— Augener,  1928,  p.  684  (part) .— Wesenberg-Lund,  1950a, 

p.  6  (part);  1950b,  p.  17  (part);  1951,  p.  10  (part). 
Eunoe  depressa  Hartman,  1948,  p.  14  (part;  not  E.  depressa  Moore,  1905). 

Description.- — Length  13-75  mm.,  width  including  setae  7-35  mm. 
Segments  36  or  37.  Prostomium  with  cephalic  peaks  short  and  blunt 
or  lacking;  anterior  pair  of  eyes  anterolateral,  visible  dorsally.  An- 
tennae with  short  papillae.  Dorsal  cirri  with  long  papillae.  Body 
flattened  ventrally,  strongly  arched  dorsally,  especially  in  anterior 
part.  With  characteristic  bulbous  structures  on  inner  sides  of  bases 
of  elytrophores  and  dorsal  tubercles.  Segmental  papUlae  quite  elon- 
gate, cylindrical,  directed  dorsally  between  the  parapodia.  Elytra 
with  fringe  of  long  papillae  on  external  border.  Ely  tral  micro  tubercles 
numerous,  close-set,  rather  low,  flattened,  semiglobose,  some  bifid; 


218  PROCEEDINGS    OF   THE    NATIONAL  MUSEUM  vol.  103 

macrotubercles  confined  mostly  to  a  single  row  near  external  border, 
dark  colored  to  pale  yellow,  nodular,  with  roughened  tips  or  a  fasicle 
of  short  spikes.  Elytra  often  covered  with  a  good  deal  of  debris  and 
foreign  material.  Color:  In  life  and  in  alcohol:  Mediodorsal 
surface  colorless  or  banded  with  olive-brown  between  elytrophores  and 
dorsal  tubercles;  ventral  surface  without  color  or  olive-brown;  setae 
yellow  or  light  amber-colored;  elytra  yellow  or  tannish  mottled  with 
reddish  brown. 

Remarks. — Eunoe  nodosa  has  been  confused  with  E.  oerstedi.  The 
two  species  have  been  separated  by  Sars  (1860),  Malmgren  (1865), 
Verrill  (1881),  Murdoch  and  Benedict  (1885,  as  Polynoe  islandica  and 
P.  scahru),  and  Treadwell  (1937).  They  have  been  considered  to  be 
synonymous  and  a  highly  variable  species  by  Theel  (1879),  Wiren 
(1883),  Fauvel  (1923),  Ditlevsen  (1917),  Augener  (1928),  and  Wesen- 
berg-Lund  (1950a,  b).  Based  on  a  study  of  the  material  from  Point 
Barrow  as  well  as  numerous  other  specimens  in  the  U.  S.  National 
Museum  from  Greenland  and  off  New  England,  the  two  species  appear 
to  be  separable  on  the  basis  of  a  number  of  characters  as  indicated 
in  the  key.  They  agree  in  having  the  prostomium  with  cephalic 
peaks  short  and  blunt  or  lacking,  the  position  of  the  eyes,  the  dorsal 
cirri  with  long  papillae,  and  the  palps  each  with  six  longitudinal  rows 
of  papillae. 

New  records. — Arctic  Alaska:  Eluitkak  Pass,  Elson  Lagoon  near 
Point  Barrow,  stony-mud;  off  Point  Barrow  base,  up  to  12.1  miles 
from  shore,  18.3-87  fathoms,  on  various  combinations  of  mud,  stones, 
pebbles,  gravel,  rocks  (14  stations,  33  specimens).  Bering  Sea:  57° 
N.,  163°  48'  W.,  38  fms.,  Alaska  King  Crab  Expedition  (Hartman 
(1948),  as  E.  depressa);  Albatross  Sta.  3252,  57°22'  N.,  164 °24'  W., 
29.5  fms.,  black  mud,  1890,  and  Sta.  3512,  57°49'  N.,  169°27'  W., 
38  fms.,  1893.  Southwestern  Alaska:  Belkofsky  Bay,  15-25  fms., 
Dall,  1880.  Franz  Josef  Land:  Aberdore  Chaimel,  east  Alger 
Island,  10  fms.,  Baldwin-Ziegler  Expedition,  1901.  East  Coast 
North  America:  Off  Labrador,  10-95  fms.,  Blue  Dolphin  Expeditions, 
1949,  1950;  Albatross  Sta.  2432,  off  Newfoundland,  43°04'  N.,  50°45' 
W.,  64  fms.,  1885;  Bay  of  Fundy,  Grand  Manan,  Nova  Scotia,  Maine, 
Massachusetts,  16-120  fms.,  U.  S.  Fish  Commission. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian,  Alaskan, 
and  Canadian  Arctic,  Baffin  Bay,  Davis  Strait,  Greenland,  Jan 
Mayen,  Spitsbergen,  Norway,  Franz  Josef  Land,  Barents  Sea,  Novaya 
Zemlya,  Kara  Sea.  Also  Bering  Sea;  north  Japan  Sea;  Iceland,  Faroes, 
Shetlands  to  English  Channel;  Hudson  Bay  to  Massachusetts.  In 
10-690  fathoms. 


MARINE    POLYCHAETE    WORMS — PETTIBONE  219 

Eunoe  oerstedi  Malmgren,  1865 

Figure  26,  d 

Lepidonote  scabra  Oersted,  1843,  p.  164,  pi.  1,  figs.  2,  7,  10,  12-13,  17-18  (not 

Aphrodita  scabra  Fabricius,  1780). 
Polynoe  scabra  Sars,  1860,  p.  58.— Hansen,  1882,  p.  11.— Wir6n,  1883,  p.  387, 

pi.  28,  fig.  1  (part).— Murdoch,  1885,  p.  152  (not  A.  scabra  Fabricius,  1780). 
Eunoe  oerstedi  Malmgren,   1865,  p.  61,  pi.  8,  fig.  3. — Moore,   1909b,  p.   135. — 

Sumner,  1913,  p.  618.— Treadwell,  1937,  p.  27.— Hartman,  1944a,  pp.  334, 

337.— Gorbunov,  1946,  p.  38.— Pettibone,  1953,  p.  46,  pi.  23. 
Eunoe  barbata  Moore,  1910,  p.  334,  pi.  28,  figs.  1-6.— Treadwell,  1925,  p.  1.— 

Hartman,  1939,  p.  53.— Berkeley  and  Berkeley,  1945,  p.  321;  1948,  p.  14, 

fig.  16.— Pettibone,  1949,  p.  2. 
Eunoe  nodosa  Fauvel,  1923,  p.  51,  fig.  18,  a-e  (part). — Hartman  and  Reish,  1950, 

p.  7  (not  Polynoe  nodosa  Sars,  1860). 
Harmothoe  nodosa  Ditlevsen,  1917,  p.  6,  pi.  2,  fig.  1  (part;  not  pi.  3,  fig.  10). — 

Augener,  1928,  p.  684  (part) .— Wesenberg-Lund,  1950a,  p.  6  (part);  1950b, 

p.  17  (part);  1951,  p.  10  (part;  not  P.  nodosa  Sars,  1860). 

Description. — Length  18-73  mm.,  width  including  setae  6-29  mm. 
Segments  37-39  (39-42  in  specimens  from  Washington).  Prostomium 
with  cephahc  peaks  poorly  developed  or  lacking;  anterior  pair  of  eyes 
anterolateral,  visible  dorsally.  Antennae  and  dorsal  cirri  with  long 
papillae.  Body  elongate,  oval  in  cross  section.  Segmental  papillae 
short  and  bulbous.  Tips  of  notosetae  may  be  worn  down  and  blunt, 
especially  in  larger  specimens.  Elytra  with  lateral  fringe  of  papillae. 
Elytral  surface  exceedingly  scabrous,  with  microtubercles  one-  to 
many-pronged;  macro  tubercles  branched,  extremely  variable  in  size, 
number,  arrangement  and  shape;  some  brownish,  some  translucent. 
Color:  In  alcohol :  Dusky,  dark  or  greenish  black  between  elytrophores 
and  dorsal  tubercles,  with  some  color  on  cirrophores  of  dorsal  cirri 
and  tips  of  parapodia;  setae  dark  amber-colored. 

Remarks. — This  species  is  closely  related  to  and  has  been  confused 
with  E.  nodosa  as  discussed  above.  Examination  of  the  type  of  E. 
barbata  Moore  from  central  California  revealed  no  essential  differences 
from  E.  oerstedi. 

New  records. — Arctic  Alaska:  Eluitkak  Pass,  Elson  Lagoon  near 
Point  Barrow,  6.6  fms.;  off  Point  Barrow  base,  up  to  8  miles  from 
shore,  18.3  to  75.5  fms.,  on  bottoms  of  mud,  sand,  shells,  pebbles, 
gravel,  rocks,  stones  (10  stations,  20  specimens).  Canadian  Arctic: 
Baffin  Island,  66°43'  N.,  80°07'  W.,  1927;  center,  south,  and  southeast 
corner  Foxe  Basin,  25-37  fms.,  1927;  Kneeland  Bay  in  Frobisher 
Bay,  Baffin  Island,  7-12  fms.,  1942;  all  collected  by  Bartlett.  West 
Greenland:  Murchison  Sound,  45-60  fms.,  Bartlett,  1938.  East 
Greenland:  Clavering  Fiord,  1939;  off  Cape  Hold  with  Hope, 
23-40  fms.,  Bartlett,  1939.  South  Greenland:  Off  Cape  Farewell, 
70  fms.,  Bartlett,  1939.     East  Coast  North  America:  Off  Labrador, 

261112—54 2 


220  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  103 

25-40  fms.,  Blue  Dolphin  Expeditions,  1949,  1950;  Nova  Scotia, 
Grand  Manan,  Maine,  Massachusetts,  low  water  to  110  fms.,  U.  S. 
Fish  Commission.  Japan:  Albatross  Sta.  3656,  Hakodate  Bay,  11.5 
fms.,  1896. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian,  Alaskan 
and  Canadian  Arctic,  Baffin  Bay,  Davis  Strait,  Greenland,  Spits- 
bergen. Also  from  Norway  to  English  Channel;  Labrador  to 
Massachusetts;  Bering  Sea  to  central  California;  and  Japan.  In 
low  water  to  516  fathoms. 

Genus  Harmothoe  Kinberg,  1857 

As  used  herein,  Harmothoe  includes  Lagisca  Malmgren,  1865; 
Evarne  Malmgren,  1865;  and  Evarnella  Chamberlin,  1919. 

Both  species  represented  have  the  body  short,  attenuated  posteriorly, 
segments  about  40  (36-50),  covered  by  elytra  except  for  posterior 
few  segments;  elytra  with  numerous  conical  micro  tubercles.  Pro- 
stomium  with  cephalic  peaks  prominent  (fig.  26,  a).  Antennae  and 
dorsal  cirri  with  short  papillae.  Notosetae  stouter  than  neurosetae, 
with  long  distal  spinous  regions  and  short,  pointed,  bare  tips. 

Key  to  the  species  of  Harmothoe  from  Point  Barrow 

1.  Anterior  pair  of  eyes  antero ventral  on  prostomium  (slightly  posterior  and 
lateral  to  cephalic  peaks),  not  visible  dorsally  (fig.  26,  a).  Elytra  with 
or  without  large,  globular  macrotubercles  near  posterior  border. 

H.  imbricata 
Anterior  pair  of  eyes  anterolateral  on  prostomium,  visible  dorsally.     Elytra 
with  macrotubercles  globular,  sausage-shaped  or  elongate,  rodlike. 

H.  extenuata 

Harmothoe  imbricata  (Linn^,  1767) 

Figure  26,  a,  e 

Aphrodita  imbricata  Linn6,  1767,  p.  1804. 

Lepidonote  cirrata  Oersted,  1843,  p.  166,  pi.  1,  figs.  1,  5-6,  11,  14-15. 

Polynoe  cirrata  Sars,  1860,  p.  60. 

Harmothoe  imbricata  Malmgren,  1865,  p.  66,  pi.  9,  fig.  8. — Webster  and  Benedict, 
1884,  p.  701;  1887,  p.  709.— Johnson,  1897,  p.  181;  1901,  p.  390.— Mcintosh, 
1900,  p.  314,  figs.— Moore,  1902,  p.  270;  1903,  p.  402;  1908,  p.  334.— Dit- 
levsen,  1909,  p.  7,  pi.  2,  fig.  5.— Sumner,  1913,  p.  617.— Southern,  1914,  p. 
52.— Chamberhn,  1920,  p.  4.— EHason,  1920,  p.  19.— Fauvel,  1923,  p.  55, 
fig.  18,  f-e;  1933,  p.  10;  1936,  p.  50.— Treadwell,  1925,  p.  1;  1937,  p.  26.— 
Augener,  1928,  p.  677;  1939,  p.  133.— Annenkova,  1934,  p.  322;  1937,  p.  151; 
1938,  p.  136.— Okuda,  1938b,  p.  83;  1939,  p.  224.— Friedrich,  1939,  p.  122.— 
Monro,  1939a,  p.  345.— Hartman,  1944a,  pp.  335,  337;  1948,  p.  13.— Thorson, 
1946,  p.  46,  figs.  17-18.— Gorbunov,  1946,  p.  38.— Zatsepin,  1948,  p.  108, 
pi.  28,  fig.  10.— Berkeley  and  Berkeley,  1948,  p.  11,  ^fig.  9.— Pettibone,  1949, 
p.  2;  1953,  p.  32,  pis.  13-16.— Hartman  and  Reish,  1950,  p.  6.— Wesenberg- 
Lund,  1950a,  p.  6;  1950b,  p.  18;  1951,  p.  12. 


MARINE    POLYCHAETE    WORMS — PETTIBONE  221 

Polynoe  imbricata  Marenzeller,  1879,  p.  9,  pi.  2,  fig.  1. — Th6el,  1879,  p.  9. — Hansen, 

1882,  pp.  11,  13,  15,  23.— Wiren,  1883,  p.  389. 
Harmothoe  levis  Treadwell,  1937,  p.  26,  figs.  1-5. 

Description. — Length  24-53  mm.,  width  includmg  setae  8-19  mm. 
(length  up  to  65  mm. — Theel,  1879).  Segments  37-39.  Prostomium 
with  anterior  pair  eyes  ventral,  slightly  posterior  and  lateral  to 
cephalic  peaks.  Elytra  with  scattered  conical  microtubercles,  with 
or  without  few  to  numerous,  small,  brownish  to  reddish,  globular  to 
elongate  cylindrical  macrotubercles  in  one,  two,  or  several  irregular 
rows  near  posterior  border;  may  be  a  few  additional  globular  papillae 
in  middle  of  elj^'tra;  with  or  without  short  elytral  fringe  of  papillae. 
Neurosetae  with  long  spinous  regions,  with  long,  bare,  hooked  tips, 
and  usually  with  a  subterminal  tooth  (may  be  broken  off).  Color: 
In  life  and  in  alcohol:  Irregularly  pigmented  middorsally,  on  cirro- 
phores,  dorsal  tubercles,  and  parapodial  lobes;  usually  without  color 
ventrally;  elytra  show  remarkable  color  variations  of  mottled  tan  and 
brown,  blackish  to  grayish,  or  reddish;  inner  halves  of  elytra  red  or 
graj'^ish  brown  resulting  in  a  middorsal,  longitudinal,  pigmented  band ; 
uniformly  tan;  tan  with  bilateral  darker  spots  near  places  of  attach- 
ment (the  color  variet}^  of  H.  levis  Treadwell) . 

Remarks. — Examination  of  the  types  of  H.  levis  Treadwell  from. 
Greenland  waters  revealed  it  to  be  but  a  color  variation  of  H. 
imbricata. 

Parasites. — Of  the  215  specimens,  13  had  parasitic  copepods, 
Herpyllobius  arcticus  Steenstrup  and  Liitken,  attached  on  the  pro- 
stomium (identified  by  Paul  L.  Illg). 

New  records. — Arctic  Alaska:  Eluitkak  Pass,  Elson  Lagoon  near 
Point  Barrow,  6.6  fms.,  gravel,  stones,  mud;  Point  Barrow  base, 
washed  ashore;  off  Point  Barrow  base,  up  to  16  miles  from  shore, 
11.7-123.5  fms.,  on  bottoms  of  stones,  mud,  masses  of  worm  tubes, 
and  various  combinations  of  mud,  pebbles,  gravel,  stones,  large 
perforated  rocks,  with  barnacles,  bryozoans,  hydroids,  tunicates, 
shells,  worm  tubes  (34  stations,  215  specimens).  Canadian  Artic: 
Southeast  corner  Foxe  Basin,  66°46'  N.,  79°15'  W.,  34-37  fms.,  1927; 
shoal  in  Kneeland  Bay,  Frobisher  Bay,  Baffin  Island,  7-12  fms., 
1942;  west  shore  Frobisher  Bay,  1942;  off  Daniels  Island,  northwest 
side  in  Newell  Sound,  Frobisher  Bay,  10-30  fms.,  1942;  all  collected 
by  Bartlett.  West  Greenland:  Oelricks  Bay,  mud,  1937;  Vaigat, 
Disko  Island,  mud,  1937;  between  Capes  Alexander  and  Chalon, 
25-40  fms.,  rocks,  1937;  Walrus  grounds,  Murchison  Sound,  45-60 
fms.,  1938;  off  Conical  Rock,  76°  N.,  67°30'  W.,  20-40  fms.,  rocks, 
shell,  1938;  all  collected  by  Bartlett.  Upernivik  Harbor,  13  fms., 
U.S.S.  Alert,  1884.  East  Greenland:  Clavering  Fiord,  1939; 
Angmogssalik,  10-15  fms.,  1939;  off  Cape  Hold  with  Hope,  23-40 
fms.,  1939;  all  collected  by  Bartlett.     Spitsbergen:  Spitsbergen  Sea, 


222  PROCEEDINGS    OF   THE    NATIONAL  MUSEUM  vol.ios 

U.S.S.  Alliance,  1881;  South  Gatt,  northwest  Spitsbergen,  79°40'  N., 
7  fms.,  E.  Wilkinson.  East  Coast  North  America:  Off  Labrador 
and  Newfoundland,  intertidal  to  60  fms.,  Blue  Dolphin  Expeditions, 
1949,  1950,  1951;  off  Nova  Scotia,  Grand  Manan,  Maine,  Massachu- 
setts, Rhode  Island,  Long  Island  Sound,  intertidal  to  110  fms.,  U.  S- 
Fish  Commission.  Bering  Sea:  Bering  Strait,  13  fms.,  Dall,  1880; 
62°54'  N.,  166°38'  W.,  22  fms.,  and  66°12'  N.,  168°54'  W.,  Stoney, 
1884;  St.  George  Island,  Pribilofs,  G.  D.  Hanna,  1914;  Albatross 
Sta.  3252,  57°22'  N.,  164°24'  W.,  29.5  fms.,  1890,  and  Sta.  3522, 
57°58'  N.,  170°09'  W.,  41  fms.,  1893.  Southwestern  Alaska:  Bay 
of  Islands,  Adak  Island,  9-16  fms.,  1873,  and  Coal  Harbor,  Unga 
Island,  1872,  Dall.  Southeastern  Alaska:  Wrangel,  W.  H.  Jones. 
Sitka,  "worm-eaten"  wood,  L.  A.  Beardslee.  Japan:  Albatross  Sta. 
3656,  Hakodate  Bay,  1896. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian,  Alaskan, 
and  Canadian  Arctic,  Davis  Strait,  Greenland,  Jan  Mayen,  Spits- 
bergen, Franz  Josef  Land,  Barents  Sea,  Novaya  Zemlya.  Also 
Iceland  and  Norway  to  Mediterranean  and  Adriatic;  Labrador  to 
New  Jersey;  Bering  Sea  to  southern  California;  and  Japan.  In  low 
water  to  2,030  fathoms. 

Harmothoe  extenuata  (Grube,  1840) 

Polyno'e  extenuata  Grube,  1840,  p.  86  {fide  Claparfede,  1868). — ClaparMe,  1868, 

p.  380,  pi.  2,  fig.  2. 
Polynoe  rarispina  Sars,  1860,  p.  60.— Th^el,  1879,  p.  8.— Hansen,  1882,  p.  7.— 

Wir^n,  1883,  p.  389. 
Lagisca  rarispina  Malmgren,  1865,  p.  65,  pi.  8,  fig.  2. — Verrill,  1881,  pp.  311, 

314.— Webster  and  Benedict,  1884,  p.  700;  1887,  p.  709.— Moore,  1902,  p. 

269;  1908,  p.  335.— Fauvel,  1911,  p.  9;  1914,  p.  65.— Chamberlin,  1920,  p. 

5.— Berkeley  and  Berkeley,  1943,  p.  129;  1948,  p.  16,  figs.  18,  19. 
Lagisca  propinqua  Malmgren,  1867,  p.  9,  pi.  1,  fig.  3,  a-e. 
Lagisca  extenuata  Marenzeller,  1876,  p.  133,  pi.  1,  fig.  1;  1904,  p.  318. — Mcintosh, 

1900,  p.  307,  figs.— Fauvel,  1923,  p.  76,  fig.  28,  a-m;  1934a,  p.  14.— Monro, 

1939a,  p.  345.— Wesenberg-Lund,  1939a,  p.  6,  fig.  2;  1950a,  p.  8;  1950b,  p. 

27;  1951,  p.  18. 
Lagisca  impatiens  Webster,  1879b,  p.  102;  1886,  p.  129,  pi.  4,  figs.  1-7. 
Polynoe  semisculpta  Hansen,   1882,  p.  26,  pi.  3,  figs.   16-20  (?P.  semisculptus 

Johnston,  1865). 
Lagisca  fioccosa  Mcintosh,  1900,  p.  298,  figs. — Southern,  1914,  p.  51  (not  Polynoe 

floccosa  Savigny,  1820). 
Harmothoe  extenuata  Alaejos  y  Sanz,  1905,  p.  55,  pi.  9,  fig.  8;  pi.  10,  figs.  1-12; 

pi.  11,  fig.  1.— Ehlers,  1913,  p.  446.— Pettibone,  1953,  p.  31. 
Harmothoe  rarispina  Ditlevsen,  1909,  p.  5,  pi.  1,  figs.  2-4;  1937,  p.  11. — Augener, 

1928,  p.  685.— Annenkova,  1937,  p.  153;  1938,  p.  136.— Friedrich,  1939,  p. 

122.— Gorbunov,  1946,  p.  38.— Zatsepin,  1948,  p.  108,  pi.  28,  fig.  15. 
Harmothoe  triannulata  Moore,  1910,  p.  346,  pi.  29,  figs.  18-22. — Hartman,  1938a, 

p.  118.— Berkeley  and  Berkeley,  1948,  p.  12,  fig.  10. 
Lagisca  extenuata  var.  spinulosa  Fauvel,  1914,  p.  64,  pi.  4,  figs.  27-29. 
Harmothoe  propinqua  Ditlevsen,  1917,  p.  14,  pi.  3,  figs.  1,  3. 
Harmothoe  rarispina  forma  propinqua  Annenkova,  1937,  p.  153;  1938,  p.  137. 


MARINE    POLYCHAETE    WORMS — PETTIBONE  223 

Evarnella  triannulata  Berkeley  and  Berkeley,  1942,  p.  188;  1943,  p.  130. — Hartman, 
1948,  p.  13  (part;  includes  H.  muliisetosa  Moore  and  H.  extenuata). — Petti- 
bone,  1949,  p.  1. 

Description. — Length  13-68  mm.,  width  including  setae  4-20  mm. 
(length  up  to  74  mm. — Ditlevsen,  1937).  Segments  37-47.  Prosto- 
mium  with  eyes  large,  anterior  pair  anterolateral,  slightly  anterior  to 
widest  part  of  prostomium,  visible  dorsally.  Neurosetae  with  en- 
larged, long  spinous  regions,  with  tips  slightly  hooked,  with  small, 
secondary  tooth  present  or  absent;  at  least  some  of  the  setae  show  a 
remnant  of  a  secondary  tooth.  The  upper  and  lower  neurosetae  tend 
to  be  unidentate  (not  always)  while  the  middle  ones  are  bidentate, 
with  a  secondary  tooth  or  remnant  of  it  (in  specimens  from  Woods 
Hole  region  only  a  few  neurosetae  have  a  secondary  tooth  or  remnant 
of  one).  Elytra  with  numerous  micro  tubercles,  more  or  less  uniform 
in  size,  conical,  mth  tips  blunt,  pointed,  or  bifid;  usually  with  few  to 
fairly  numerous  macrotubercles  distinctly  set  off  from  elytral  surface, 
usually  narrower  at  the  base,  brown,  smooth,  globular,  sausage- 
shaped,  or  elongate  fusiform,  variable  in  number,  0-9  near  posterior 
border,  0-13  scattered  near  center  of  elytron.  Elytra  with  short 
fringe  of  papillae  on  external  border.  Color:  In  alcohol:  Without 
color  anteriorly;  on  posterior  half,  with  brownish  transverse  bands  and 
two  brown  spots  basallj''  on  cirrophores  (in  specimens  from  Woods 
Hole  region,  found  intertidally,  the  body  is  darldy  pigmented  grayish- 
green  middorsally) ;  antennae  and  dorsal  cirri  ringed  with  brown; 
elytra  mottled  with  brown,  sometimes  with  a  darker  spot  posterior 
and  medial  to  the  place  of  attachment;  the  micro  tubercles  may  be 
covered  with  a  brown  extraneous  material,  giving  a  streaked  appearance. 

Remarks. — H.  extenuata  is  an  extremely  variable  species  as  indicated 
by  the  number  of  names  that  have  been  applied  to  it.  It  reaches  its 
greatest  size  in  arctic  waters;  it  is  small  when  found  intertidally,  as 
in  the  Puget  Sound  region  (up  to  23  mm. — H.  triannulata  of  Moore). 
When  found  intertidally  off  New  England,  it  also  is  small  (about  26 
mm.),  and  the  secondary  tooth  of  the  neurosetae  is  suppressed,  usually 
with  only  a  remnant  showing  on  a  few  neurosetae,  the  rest  being  entire 
(L.  impatiens  of  Webster).  In  the  collection  from  Point  Barrow, 
where  it  was  the  most  abundant  polychaete,  it  is  represented  by 
specimens  of  all  sizes  from  very  small  to  large  (up  to  68  mm.). 

Parasites. — Of  the  375  specimens,  3  had  the  parasitic  copepod 
Herpyllohius  arcticus  Steenstrup  and  Lutken  attached  to  the  pros- 
tomium (identified  by  lUg) . 

New  records. — Arctic  Alaska:  Eluitkak  Pass,  Elson  Lagoon  near 
Point  Barrow,  6.6  fms.;  off  Point  Barrow  base,  up  to  15  miles  from 
shore,  13.3-123.5  fms.,  on  bottoms  of  mud,  stones,  masses  of  worm 
tubes,  and  various  combinations  of  mud,  pebbles,  rocks,  gravel, 
stones,    large   perforated   rocks,    with   shells,    bryozoans,   hydroids, 


224  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  103 

worm  tubes,  from  starfish  and  crab,  Hyas  coarctatus  (37  stations,  375 
specimens).  Canadian  Arctic:  Foxe  Basin,  66°30'  N.,  80°  W., 
1927;  center  Foxe  Basin,  25-31  fms.,  1927;  7  miles  east  Cape  Dor- 
chester, Foxe  Channel,  25  fms.,  1927;  Baffin  Island,  66°43'  N.,  80°07' 
W.,  1927;  Hurd  Channel  between  Bushman  Island  and  Melville 
Peninsula,  11  fms.,  rocky,  1933;  south  end  Cape  Martineau,  Melville 
Peninsula,  7-15  fms.,  1933;  3  miles  south  Salisbury  Island,  Hudson 
Strait,  27  fms.,  1933;  south  end  Cobourg  Island,  Baffin  Bay,  75°40' 
N.,  78°58'  W.,  11.3-20  fms.,  1935;  east  end  Cobourg  Island,  75°40' 
N.,  78°50'  W.,  23-36  fms.,  1935;  southeast  corner  Foxe  Basm, 
66°46'  N.,  79°15'  W.,  34-37  fms.,  1937;  Kneeland  Bay,  Frobisher 
Bay,  Baffin  Island,  7-12  fms.,  1942;  all  collected  by  Bartlett.  West 
Greenland:  Vaigat,  Disko  Island,  mud,  1937;  between  Capes 
Alexander  and  Chalon,  25-40  fms.,  rocky,  1937;  Walrus  grounds, 
Murchison  Sound,  45  fms.,  1938;  all  collected  by  Bartlett.  North 
Greenland:  North  Omenolu  near  North  Star  Bay,  17  fms.,  and  Cape 
Alexander,  entrance  to  Smith  Sound,  rocky,  1932,  Bartlett.  East 
Greenland:  Off  Cape  Hold  with  Hope,  23-40  fms.,  Bartlett,  1939. 
South  Greenland:  Cape  Farewell,  70  fms.,  Bartlett,  1939.  Franz 
Josef  Land:  Aberdore  Channel  east  Alger  Island,  10  fms.,  Baldwin- 
Ziegler  Polar  Expedition,  1901.  Bering  Sea:  Alaska  King  Crab 
Investigation,  1941,  Sta.  D8-41,  58°34'  N.,  165°17'  W.,  42  fms.; 
Sta.  D7-41,  57°  N.,  163°48'  W.,  38  fms.;  Sta.  Dll-41,  12  miles  east 
Walrus  Island,  Pribilofs,  31-33  fms.;  St.  George  Island,  PribUofs, 
village  beach,  1914,  and  St.  Paul  Island,  Pribilofs,  1915,  Hanna; 
62°54'  N.,  166°38'  W.,  22  fms.,  and  62°15'  N.,  167°48'  W.,  20  fms., 
Stoney,  1884.  Southwestern  Alaska:  Alaska  King  Crab  Investiga- 
tion, 1940,  1941,  Sta.:  89-40,  Unga  Strait,  37-47  fms.;  93-40,  Spitz 
Island,  55-68  fms.;  Sand  Point;  128-40,  Shelikof  Strait  off  Hallo  Bay, 
35-48  fms.;  60-40,  Lenard  Harbor,  20-25  fms.;  L18-41,  Kupreanof 
Strait,  south  side,  2  miles  northwest  Bare  Island,  13-15  fms.;  72-40, 
Cold  Bay,  15-50  fms.;  52-40,  Canoe  Bay,  35-40  fms.,  on  Paralithodes 
camtschatica;  100-40,  Alitak  Bay,  30  fms.;  59-40,  between  inner 
Iliasik  and  Goloi  Island,  20-30  fms.  Southeastern  Alaska: 
Lituya  Bay,  6-9  fms.,  Dall,  1874.  East  Coast  North  America: 
Off  Labrador,  6-100  fms..  Blue  Dolphin  Expeditions,  1949,  1950, 
1951;  Bay  of  Fundy,  Gulf  of  Maine,  Massachusetts,  Long  Island 
Sound,  4-134  fms.,  U.  S.  Fish  Commission;  Woods  Hole  region, 
intertidal  and  dredged,  Pettibone,  1950,  1951;  Fish  Hawk  Sta.  8826, 
8898,  Chesapeake  Bay,  1920. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian,  Alaskan, 
and  Canadian  Ai-ctic,  Baffin  Bay,  Davis  Strait,  Greenland,  Spits- 
bergen, Franz  Josef  Land,  Novaya  Zemlya,  Kara  Sea.  Also  Iceland, 
Faroes,  Norway  to  Mediterranean  and  Adriatic;  Hudson  Bay  to 
Chesapeake  Bay;  Bering  Sea  to  southern  California;  north  Japan  Sea; 
South  Africa.     In  low  water  to  1,000  fathoms. 


MARINE    POLYCHAETE    WORMS — PETTIBONE  225 

G^nus  Enipo  Malmgren,  1865,  sensu  Levinsen,  1883 
Enipo  gracilis  Verrill,  1874 

Enipo  gracilis  VerriU,  1874a,  pp.  407,  411,  pi.  6,  fig.  4;  1874b,  pp.  347,  361,  378, 

pi.  5,  fig.  3.— Pettibone,  1953,  p.  22,  pi.  7. 
Enipo  cirrata  Treadwell,  1925,  p.  1,  figs.  1-4. — Hartman,  1938a,  p.  120,  &g.  39, 

d,  e.— Berkeley  and  Berkeley,  1942,  p.  188;  1945,  p.  322;  1948,  p.  16,  fig.  20. 
Polynoe  gracilis  Hartman,  1942a,  p.  26,  figs.  23-26, 

Description. — Length  25-76  mm.,  width  including  setae  3-4  mm. 
Segments  variable  in  number  (45,  with  regenerating  posterior  end, 
to  74).  Body  elongate,  slender,  with  sides  nearly  parallel.  Prosto- 
mium  without  cephalic  peaks.  In  some  specimens  from  New  England 
there  are  slight  prostomial  peaks  (prostomial  peaks  present — Hart- 
man, 1942).  Antennae  and  dorsal  cirri  with  short,  clavate  papillae. 
Dorsal  cirri  short,  barely  reaching  tips  of  setae  in  anterior  part,  longer 
in  posterior  region  without  elytra.  Elytra  oval,  translucent,  small, 
usually  leaving  middorsum  uncovered,  smooth,  without  papillae  or 
tubercles.  Notosetae  form  a  short,  bushy  bundle,  rather  numerous 
(more  than  30  in  specimens  from  Point  Barrow)  to  few  (2-20  or 
rarely  0 — Treadwell,  1925),  slender  to  moderately  stout,  tapering 
gradually  to  short,  bare,  blunt  tips,  with  spinous  rows  distally. 
Neurosetae  stouter  than  notosetae,  upper  ones  with  longer  spinous 
regions  (17-23  or  so  rows),  middle  ones  with  shorter  spinous  regions 
(10  or  so  rows),  with  long,  bare,  slightly  hooked  tips  (neurosetae  may 
be  all  similar,  with  5-10  transverse  spinous  rows) .  Color  :  In  alcohol : 
Without  color  or  brownish  middorsally;  elytra  pigmented  smoky 
brown  on  the  medial  halves. 

Remarks. — -Known  to  be  commensal  with  the  maldanid  Nicomache 
lumhricalis  (Fabricius)  in  Alaska  (as  E.  cirrata — Berkeley  and  Berke- 
ley, 1942)  and  off  Halifax,  Nova  Scotia,  and  Cape  Cod  (specimens  in 
the  U.  S.  National  Museum,  identified  by  Verrill). 

New  Records. — Arctic  Alaska:  Off  Point  Barrow,  up  to  12.1  miles 
from  shore,  21-123.5  fms.,  from  breaking  rock,  masses  of  worm  tubes, 
with  stones,  large  perforated  rocks  (3  stations,  3  specimens).  East 
Coast  North  America:  Off  Nova  Scotia,  Maine,  Massachusetts,  2- 
198  fms.,  U.  S.  Fish  Commission. 

Distribution. — Alaskan  Arctic;  southwestern  Alaska  to  Washington; 
Nova  Scotia  to  Massachusetts.     In  2-123.5  fathoms. 

Genus  Arcteobia  Annenkova,  1934 
Arcteobia  anticostiensis  (Mcintosh,  1874) 

Eupolynoe  anticostiensis  Mcintosh,  1874,  p.  265,  pi.  10,  figs.  1-4. — Wir6n,  1883, 

p.  390,  pi.  29,  fig.  2.— Marenzeller,  1890,  pi.  1. 
Harmothoe  anticostiensis  Augener,  1928,  p.  691,  pi.  11,  fig.  13. 


226  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  los 

Ardeobia  anticostiensis  Annenkova,  1934,  p.  322;  1937,  p.  149,  pi.  3,  figs.  26,  27; 

1938,  p.  133. 
Eucranta  anticostiensis  Hartman,  1944a,  p.  337. 

Description. — Length  11-26  mm.,  width  mcluding  setae  4-8  mm. 
Segments  35  or  36.  Prostomium  with  distinct  cephaHc  peaks;  anterior 
pair  of  eyes  antero ventral,  not  visible  dorsally.  Antennae  and  dorsal 
cirri  with  short,  clavate  papillae.  Elytral  pairs  get  larger  posteriorly, 
the  last  pair  usually  extending  beyond  end  of  body.  Elytra  without 
fringe  of  papillae,  smooth  except  for  scattered  microtubercles  on 
anterior  curved  part.  Upper  notosetae  shorter,  stouter,  with  short, 
blunt  to  sharp-pointed  tips;  most  of  notosetae  more  slender,  with 
capillary  tips.  Few  upper  neurosetae  longer,  ending  in  sharp,  slender 
tips;mostofnem'Osetae  with  bifid  tips.  Color:  In  life  and  iu  alcohol : 
Irregularly  banded  middorsally,  gi-eenish  to  greenish  black,  with  a 
transverse  stripe  and  often  a  pair  of  small  spots;  elytra  with  reddish 
brown  pigmented  area,  usually  C-shaped  on  inner  and  posterior  parts 
and  often  a  darker  spot  over  place  of  attachment;  may  form  a  com- 
plete, circular,  colored  area. 

Remarks.- — -Probably  commensal  in  habit;  off  Labrador,  found  com- 
mensal in  the  sinuous  tubes  of  the  terebeUid  Pista  Jlexuosa  (Grube), 
one  worm  per  tube. 

New  records. — Arctic  Alaska:  Point  Barrow  base,  washed  ashore; 
off  Pomt  Barrow  base,  up  to  12.1  miles  from  shore,  24.7-123.5  fms., 
on  bottoms  of  mud,  masses  of  worm  tubes,  and  various  combinations 
of  mud,  gravel,  stones,  rocks,  and  large  perforated  rocks  (7  stations, 
14  specimens).  East  Coast  North  America:  Off  Labrador,  40-95 
fms..  Blue  Dolphin  Expedition,  1949;  off  Nova  Scotia,  Maine,  Mas- 
sachusetts, 10-86  fms.,  U.  S.  Fish  Commission. 

Distribution. — Alaskan  Arctic;  Labrador  to  Massachusetts;  Bering 
Sea;  north  Japan  Sea.     In  low  water  to  123.5  fathoms. 

Genus  Gattyana  IVIcIntosh,  1897 

Key  to  the  species  of  Gattyana  from  Point  Barrow 

1.  Notosetae  with  tips  blunt  to  capillary.  Anterior  pair  of  eyes  anteroventral, 
not  visible  dorsally;  cephalic  peaks  distinct.  Elytra  with  microtubercles 
only G.  cirrosa 

AH  notosetae  with  capillary  tips.  Anterior  pair  of  eyes  anterolateral;  cephalic 
peaks  short  and  blunt.  Elytra  with  conical  and  bifid  microtubercles,  with  a 
few  larger,  knobUke  macrotubercles G.  ciliata 

Gattyana  cirrosa  (Pallas,  1766) 

Figure  26,  b 

Aphrodita  cirrhosa  Pallas,  1766,  p.  95,  pi.  8,  figs.  3-6. 

Nychia  cirrosa  Malmgren,  1865,  p.  58,  pi.  8,  fig.  1.— Th6el,  1879,  p.  7.— VerriU, 

1881,  pp.  306,  311.— Wir6n,  1883,  p.  387.— Webster  and  Benedict,  1884,  p. 

700;  1887,  p.  708.—  ?  Andrews,  1891,  p.  279. 


MARINE    POLYCHAETE    WORMS — PETTIBONE  227 

Gattyana  cirrosa  Mcintosh,  1900,  p.  285,  figs.— Moore,  1902,  p.  259;  1908,  p. 
337.— Southern,  1914,  p.  51.— EHason,  1920,  p.  21.— Fauvel,  1923,  p.  49, 
fig.  17,  a-g.— Augener,  1928,  p.  692;  1939,  p.  136.— Gustafson,  1936,  p.  5.— 
TreadweU,  1937,  p.  25.— Annenkova,  1937,  p.  148;  1938,  pp.  83,  96,  133, 
222.— Monro,  1939a,  p.  345.— Berkeley  and  Berkeley,  1943,  p.  129;  1948, 
p.  13,  fig.  13.— Thorson,  1946,  p.  45.— Gorbunov,  1946,  p.  38.— Hartman, 
1948,  p.  14.— Zatsepin,  1948,  p.  106,  pi.  28,  fig.  5.— Pettibone,  1949,  p.  2; 
1953,  p.  41,  pi.  20.— Wesenberg-Lund,  1950a,  p.  5,  pi.  1,  figs.  1,  2;  1950b, 
p.  12;  1951,  p.  9. 

Gattyana  cirrhosa  Chamberlin,  1920,  p.  8. 

Description. — Length  11-35  mm.,  width  including  setae  4-12  mm- 
Segments  35-38.  Elytra  with  long  papillae  on  posterior  and  external 
borders  as  well  as  scattered  over  the  exposed  part  of  elytral  surface, 
usually  covered  with  debris,  giving  a  straggly  appearance.  Elytral 
tubercles  amber-colored,  exceedingly  variable  in  shape,  conical  or 
cylindrical,  simple,  bifid  or  quadrifid,  showing  various  degrees  of 
being  worn  down;  tubercles  on  the  anterior  few  pairs  of  elytra  tend 
to  be  less  worn  down,  showing  their  more  characteristic  shapes. 
Color:  In  alcohol:  Elytra  tan  mottled  with  brown  or  uniformly  tan; 
often  with  blackish  foreign  material  on  elytral  tubercles  and  papillae. 
Specimens  washed  ashore  were  essentially  free  from  debris,  the  elytra 
whitish,  iridescent,  with  amber-colored  tubercles  and  inconspicuous, 
delicate  papillae. 

Parasites. — Three  of  the  126  specimens  had  the  parasitic  copepod 
Herpyllobius  arcticus  Steenstrup  and  Liitken  attached  to  the  prostomia 
(identified  by  Illg). 

New  records. — Arctic  Alaska:  Eluitkak  Pass,  Elson  Lagoon  near 
Point  Barrow,  6.6  fms.;  Point  Barrow  base,  washed  ashore;  off  Point 
Barrow  base,  up  to  15  miles  from  shore,  18.3-123.5  fms.,  on  bottoms 
of  mud,  stones,  masses  of  worm  tubes,  and  various  combinations  of 
mud,  pebbles,  small  and  large  rocks,  stones,  gravel,  and  worm  tubes; 
from  breaking  rock  (30  stations,  126  specimens).  Cape  Smyth,  2.5-3 
fms..  Point  Barrow  Expedition,  1883.  Canadian  Arctic:  South  end 
Cape  Martineau,  Melville  Peninsula,  7-15  fms.,  1933;  Foxe  Basin, 
1933;  east  end  Cobourg  Island,  Baffin  Bay,  75°40'  N.,  78°50'  W., 
23-40  fms.,  1935;  Kneeland  Bay,  Frobisher  Bay,  Baffin  Island,  14 
fms.,  1942;  all  collected  by  Bartlett.  West  Greenland:  Vaigat, 
Disko  Island,  1937;  between  Capes  Alexander  and  Chalon,  25-40  fms., 
rocky,  1937;  Walrus  grounds,  Murchison  Sound,  60  fms.,  1938; 
northwest  Conical  Rock,  1938;  all  collected  by  Bartlett.  Upernivik 
Harbor,  13  fms.,  and  Godhavn,  U.  S.  S.  Alert,  1884.  East  Green- 
land: Off  Cape  Hold  with  Hope,  4-6  fms.,  Bartlett,  1939.  North 
Greenland:  North  Omenolu  near  North  Star  Bay,  17  fms.,  Bartlett, 
1932.  SpnsBERGEN:  Spitsbergen  Sea,  U.  S.  S.  ylZ^iance,  1881.  Bering 
Sea:  Bering  Strait,  13  fms.,  Dall,  1880.    Albatross  Sta.  3326,  53°40' 


228  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  tol.  103 

N.,  167°41'  W.,  1890.  Southwestern  Alaska:  Iluiliiik  Harbor, 
Unalaska,  Dall,  1871.  Southeastern  Alaska:  Sitka,  Dall.  East 
Coast  North  America:  Off  Labrador,  13-125  fms.,  Blue  Dolphin 
Expeditions,  1949,  1950,  1951. 

Distribution. — -Widely  distributed  in  the  Arctic:  Siberian,  Alaskan, 
and  Canadian  Arctic,  Davis  Strait,  Greenland,  Spitsbergen,  Novaya 
Zemlya.  Also  Iceland,  Faroes,  Norway  to  France;  Hudson  Bay  to 
Massachusetts;  Bering  Sea  to  Washington;  north  Japan  Sea.  In 
low  water  to  630  fathoms. 

Gattyana  ciliata  Moore,  1902 

Gattyana  ciliata  Moore,  1902,  p.  263,  pi.  13,  figs.  14-19;  pi.  14,  fig.  20;  1905a, 
p.  525;  1908,  p.  337.— Annenkova,  1937,  p.  148,  pi.  2,  figs.  9,  11;  pi.  4,  fig.  33; 
1938,  pp.  83,  100,  132,  224.— Berkeley  and  Berkeley,  1948,  p.  12,  fig.  11.— 
Pettibone,  1953,  p.  40,  pi.  19. 

Description.- — Length  63-65  mm.,  width  including  setae  25-29  mm, 
(length  up  to  80  mm.^ — ^Annenkova,  1937).  Segments  36  or  37. 
Cephalic  peaks  lacking  or  short  and  blunt.  Elytral  tubercles  of 
several  kinds:  Microtubercles  numerous,  conical,  pointed,  hooked, 
some  bifid;  usually  intermediate-sized  tubercles  elongated,  conical, 
some  bifid,  in  several  rows  arranged  diagonally  from  center  of  elytron 
lateralljT^;  conical  macro  tubercles  Vv^ith  blunt,  roughened  tips,  usually 
near  posterior  border  of  elytra,  variable  in  number,  size,  and  shape. 
Color:  In  alcohol:  Elytra  tannish  mottled  with  brown. 

New  records. — Arctic  Alaska:  Oft'  Point  Barrow  base,  up  to  12.1 
miles  from  shore,  20-87  fms.,  on  bottoms  of  rocks,  stones,  smaU 
amount  of  gravel  (4  stations,  4  specimens).  Bering  Sea:  Albatross 
Sta.  3606,  55°27'  N.,  167°47'  W.,  87  fms.,  green  mud,  fine  sand,  1895. 

Distribution. — Alaskan  Arctic  (originally  recorded  from  Greenland 
waters,  corrected  to  Icy  Cape,  Alaska — Moore,  1902,  1905);  Bering 
Sea  to  Washington;  north  Japan  Sea.     In  4.4-303  fathoms. 

Family  Sigalionidae 

Prostomium  subglobular,  with  two  pairs  of  eyes,  dorsal  antennae 
1-3,  a  pair  of  subulate,  ventral  palps  (fig.  26,/).  First  or  tentacular 
segment  with  or  without  numerous  setae  (lacldng  in  Pholoe),  with 
two  pairs  of  tentacular  cirri.  Parapodia  biramous,  with  notosetae 
simple,  neurosetae  simple  or  composite  or  both;  with  paired  dorsal 
elytra  on  certain  segments ;  without  dorsal  cirri ;  with  paired  subulate 
ventral  cirri.  Pygidium  with  pair  of  anal  cirri.  With  or  without 
cirriform  branchiae  and  ciliated  cushions  or  ctenidia  on  parapodia 
(both  lacking  in  Pholoe) .  Muscular  proboscis  eversible,  with  circle  of 
soft  marginal  papillae  and  two  pairs  of  interlocking,  chitinous  jaws. 

Represented  by  a  single  species  from  Point  Barrow. 


MARINE    POLYCHAETE    WORMS — PETTIBONE 


Figure  26. — Polynoidae:  a,  Harmothoe  imbricata,  dorsal  view  anterior  end,  first  pair  elytra 
removed;  h,  Gattyana  cirrosa,  part  of  elytron  enlarged,  showing  elytral  tubercles  and 
papillae;  c,  Euno'e  nodosa,  dorsal  view  part  of  two  segments  showing  extra  rounded  lobes 
on  inner  sides  of  elytrophore  and  dorsal  tubercle;  d,  Eunoe  oerstedi,  biramous  parapodium; 
Cy  Harmothoe  imbricata,  bidentate  neuroseta.  Sigalionidae;  /,  Pholoe  minuta,  dorsal 
view  anterior  end,  first  pair  elytra  removed.     (For  explanation  of  symbols,  see  p.  210.) 


230  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  m 

Genus  Pholoe  Johnston,  1839 
Pholoe  minuta  (Fabric! us,  1780) 

Figure  26,  / 

Aphrodita  minuta  Fabricius,  1780,  p.  314. 

Aphrodila  longa  Fabricius,  1780,  p.  313. 

Pholoe  minuta  Oersted,  1843,  p.  169,  pi.  1,  figs.  3,  4,  8,  9,  16. — Malmgren,  1865, 
p.  89,  pi.  11,  fig.  13.— Th6el,  1879,  p.  24.— Webster  and  Benedict,  1884, 
p.  701;  1887,  p.  709.— Mcintosh,  1900,  p.  437,  figs.— Moore,  1902,  p.  274; 
1908,  p.  338. — Ehlers,  1913,  p.  450.— Southern,  1914,  p.  57. — Chamberlin, 
1920,  p.  5.— Eliason,  1920,  p.  22.— Fauvel,  1923,  p.  120,  fig.  44,  a-h.— 
Augener,  1928,  p.  673.— Annenkova,  1934,  p.  322;  1937,  p.  155;  1938,  p.  138.— 
Berkeley  and  Berkeley,  1942,  p.  189.— Hartman,  1944a,  pp.  335,  337;  1948, 
p.  15.— Thorson,  1946,  p.  49,  fig.  19;  Gorbunov,  1946,  p.  38.— Zatsepin, 
1948,  p.  109,  pi.  28,  fig.  17. — Hartman  and  Reish,  1950,  p.  8.— Wesenberg- 
Lund,  1950a,  p.  9;  1950b,  p.  31;  1951,  p.  22.— Pettibone,  1953,  p.  77,  pi.  39. 

Pholoe  tuberculata  Southern,  1914,  p.  57,  pi.  6,  fig.  14,  A-L. — Berkeley  and 
Berkeley,  1945,  p.  323;  1948,  p.  22,  fig.  26. 

Description. — Length  5-25  mm.,  width  including  setae  1-4  mm. 
Segments  36-84.  Body  small,  elongate,  nearly  linear,  flattened 
dorsoventrally;  fragments  easily.  Lateral  ventral  surface  and  ventral 
sides  of  parapodial  lobes  thickly  set  with  small,  globular  to  elongate 
papillae,  usually  covered  with  debris.  Prostomium  small,  oval; 
anterior  and  posterior  pairs  of  eyes  closely  approximated.  Single 
median  antenna  short,  subulate  (lateral  antennae  lacking).  Ten- 
tacular segment  achaetous;  tentacular  cirri  similar  to  median  antenna. 
A  digitiform  facial  tubercle  on  a  rounded  lobe  dorsal  to  the  mouth 
and  ventral  to  the  prostomium.  (On  this  character,  Southern  (1914) 
differentiated  P.  tuberculata;  the  lobe,  on  which  the  facial  tubercle  is 
located,  appears  to  be  retractile;  surrounded  by  the  bases  of  the  palps, 
tentacular  cirri  and  median  antenna,  the  facial  tubercle  is  easily 
overlooked  unless  the  region  is  quite  relaxed.)  Elytra  numerous,  on 
segments  2,  4,  5,  7,  9,  11,  13,  15,  17,  19,  21,  23,  24,  continuing  on  all 
segments  to  end  of  body;  elytra  nearly  cover  the  dorsum  except  for  a 
narrow  middorsal  part,  usually  pressed  close  to  body;  elytral  surface 
smooth,  with  few  microtubercles,  somewhat  areolate  as  seen  under 
magnification,  posterior  border  with  few  scattered  papillae,  somewhat 
moniliform.  Notosetae  simple,  slender,  finely  spinous,  tapering  to 
capillary  tips,  upper  ones  shorter,  with  pronounced  angles,  the  rest 
longer,  with  more  gradual  curves.  Neurosetae  stouter  than  notosetae, 
compound,  with  short,  falcate  terminal  pieces.  Color:  In  alcohol: 
Without  color  or  irregularly  pigmented  greenish  gray;  elytra  trans- 
lucent, colorless,  or  greenish  gray  with  a  light  spot  over  the  place  of 
attachment,  or  brownish;  often  with  much  debris. 


MARINE    POLYCHAETE    WORMS — PETTIBONE  231 

Nev)  records. — Arctic  Alaska:  Eluitkak  Pass,  Elson  Lagoon  near 
Point  Barrow,  6.6  fms.;  off  Point  Barrow  base,  up  to  12.1  miles  from 
shore,  16.7-123.5  fms.,  on  bottoms  of  stones,  masses  of  worm  tubes, 
and  various  combinations  of  mud,  gravel,  stones,  rocks,  large  per- 
forated rocks,  with  bryozoans,  hydroids;  from  screen  trap  through 
hole  in  ice,  mud  (16  stations,  30  specimens).  West  Greenland: 
Vaigat,  Disko  Island,  mud,  1937,  and  Walrus  grounds,  Murchison 
Sound,  45  fms.,  1938,  Bartlett.  East  Coast  North  America:  Off 
Labrador,  6  fms..  Blue  Dolphin  Expedition,  1949;  off  Nova  Scotia, 
St.  Georges  Bank,  Maine,  Massachusetts,  Rhode  Island,  intertidal  to 
110  fms.,  U.  S.  Fish  Commission. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian,  Alaskan, 
and  Canadian  Arctic,  Davis  Strait,  Greenland,  Jan  Mayen,  Norwegian 
Sea,  Spitsbergen,  Franz  Josef  Land,  Novaya  Zemlya,  Kara  Sea. 
Also  Iceland,  Faroes,  Shetlands,  Norway  to  northern  France ;  Labrador 
to  Rhode  Island;  Bering  Sea  to  southern  Oregon;  north  Japan  Sea; 
off  South  Africa.     In  low  water  to  1,254  fathoms. 

Family  Phyllodocidae 

Prostomium  subcorneal,  suboval  or  cordiform,  with  two  eyes,  four 
or  five  antennae,  without  palps  (fig.  27,  a-<i).  Anterior  segments 
1-3  modified,  with  two  to  four  pairs  tentacular  cirri.  Parapodia 
uniramous  (exceptionally  biramous);  setae  compound  (some  may  be 
simple).  Dorsal  and  ventral  cirri  foliaceous  or  more  or  less  globular 
(fig.  27,  e-4).  Two  anal  cirri.  Proboscis  eversible,  with  papillae, 
unarmed.     Active;  mucus  secreted  in  quantities. 

Represented  by  four  genera  and  seven  species.  All  genera  repre- 
sented have  uniramous  parapodia,  with  compound  setae. 

Key  to  the  genera  of  PhyUodocidae  from  Point  Barrow 

1.  Tentacular  cirri  4  pairs;  first  tentacular  segment  rudimentary,  not  visible 
dorsally,  with  1  pair  tentacular  cirri  lateral  to  prostomium;  second  segment 
distinct,  with  2  pairs  tentacular  cirri;  third  segment  distinct,  with  1  pair 
tentacular  cirri,  1  pair  normal  ventral  cirri  (fig.  27,  c,  d) 2 

Tentacular  cirri  3  pairs;  first  tentacular  segment  distinct  dorsally,  with  1  pair 
tentacular  cirri;  second  segment  distinct,  with  2  pairs  tentacular  cirri  (fig. 
27,  b).     Prostomium  oval,  without  occipital  tubercle Mystides  (p.  232) 

Tentacular  cirri  2  pairs  on  first  achaetous  segment  (fig.  27,  a) .  Without  dorsal 
cirri  on  second  segment  which  has  a  setigerous  lobe  (may  be  very  small  or 
lacking)  and  a  foliaceous  ventral  cirrus.  Prostomium  somewhat  triangular, 
widest  basally,  with  anterior  part  rounded,  with  2  pairs  short,  subulate, 
frontal  antennae,  usually  with  2  deep-set  eyes  (may  not  be  visible  when 
preserved),  usually  with  a  median  occipital  depression  containing  a  small 
occipital  tubercle  (not  always  conspicuous),  with  a  pair  of  lateral  nuchal 
grooves Eteone  (p.  232) 


232  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  103 

2.  Antennae  4  (2  pairs  frontal  antennae).  Prostomium  cordiform,  with  an 
occipital  tubercle  in  the  posterior  notch  (fig.  27,  d) .  Proboscis  with  papillae 
proximally  arranged  in  longitudinal  rows Phyllodoce  (p.  236) 

Antennae  5  (2  pairs  frontal  antennae  and  unpaired  median  antenna) .  Prosto- 
mium oval  or  bluntly  conical,  without  occipital  tubercle  (fig.  27,  c).  Pro- 
boscis with  surface  smooth,  wrinkled,  or  with  few  scattered  papillae 

Eumida  (p.  238) 

Genus  My  slides  Theel,  1879 
Mystides  borealis  Theel,  1879 

Figure  27,  b 

Mystides  borealis  Thdel,  1879,  p.  35,  pi.  2,  figs.  29-32.— Southern,  1914,  p.  72, 
pi.  8,  fig.  19,  A-D.— Bergstrom,  1914,  p.  176,  fig.  64.— Fauvel,  1923,  p.  181, 
fig.  65,  a-d.— Augener,  1928,  p.  711.— Zatsepin,  1948,  p.  113,  pi.  29,  fig.  15.— 
Wesenberg-Lund,  1950b,  p.  37. 

Mystides  viridis  Webster  and  Benedict,  1887,  p.  712,  pi.  1,  figs.  10,  11,  13;  pi.  2, 
fig.  12.  (Type  in  USNM.) 

Mystides  notialis  Ehlers,  1913,  p.  457,  pi.  29,  figs.  1-4. 

Description. — Length  5-9  mm.,  width  0.5  mm.  Segments  33-64 
(up  to  16  mm.  long  and  73  segments — Ehlers,  1913).  Body  small, 
linear,  tapering  anteriorly  and  posteriorly,  flattened  dorsoventrally. 
Prostomium  ova],  wider  than  long,  with  two  deeply-set  eyes  in  middle 
of  prostomium,  with  four  long,  filiform  antennae  about  one-third 
way  back  from  anterior  tip.  Tentacular  cirri  3  pairs,  enlarged  basally, 
with  long,  filiform  tips.  Dorsal,  ventral,  and  anal  cirri  oval,  thick, 
flattened.  Color:  In  alcohol:  Without  color  or  irregularly  pigmented 
with  brown;  cirri  deep  reddish  brown. 

Remarks. — The  description  of  M.  notialis  from  the  Antarctic  by 
Ehlers  (1913)  follows  closely  that  of  M.  borealis  and  is  herein  considered 
to  be  synonymous. 

New  records. — ^Arctic  Alaska:  Off  Point  Barrow,  7.5  miles  from 
shore,  36  fms.,  on  bottom  of  stones,  perforated  rocks  (1  station,  2 
specimens).  West  Coast  North  America:  Off  Flat  Point,  Lopez 
Island,  Washington  Sound,  Pettibone  (1  specimen). 

Distribution. — Scattered  records  in  the  Arctic:  Arctic  Alaska,  West 
Greenland,  Spitsbergen,  Novaya  Zemlya.  Also  Ireland,  Madeira, 
Mediterranean;  east  coast  North  America  (Maine);  west  coast  North 
America  (Washington  Sound);  Antarctic  (Kerguelen).  In  4-214 
fathoms. 

Genus  Eteone  Savigny,  1817 

The  four  species  represented  have  the  body  elongate,  somewhat 
flattened  dorsoventrall}^,  tapering  anteriorly  and  posteriorly,  with 
segments  numerous  (100-300).  Proboscis  with  a  circle  of  soft  papillae 
around  opening. 


MARINE    POLYCHAETE    WORMS — PETTIBONE  233 

Key  to  the  species  of  Eteone  from  Point  Barrow 

1.  Proboscis  with  2  lateral,  longitudinal  rows  of  numerous  soft  papillae.     Dorsal 

pair  tentacular  cirri  slightly  longer  or  up  to  2-3  times  longer  than  ventral 
pair.  Dorsal  cirri  wider  than  long,  nearly  symmetrical,  subcircular  or 
slightly  lanceolate  (fig.  27,  e).  Color  in  alcohol:  Darkly  pigmented  with 
bluish-violet  iridescence,  with  longitudinal,  lighter  colored  bands  on  each 
side  of  median  dorsal  line  (or  3  dorsal  longitudinal  dark  bands — 1  median 
and   2   lateral).     Anal   cirri   subcylindrical,    3-4   times   longer   than   wide. 

Second  segment  with  setigerous  lobe  and  setae  well  developed E.  barbata 

Surface  of  proboscis  smooth,  rugose  or  irregularly  papillate.     Two  pairs  ten- 
tacular cirri  subequal  or  ventral  pair  somewhat  longer  than  dorsal  pair —  2 

2.  Dorsal  cirri  small,  not  much  larger  than  parapodial  lobes,  longer  than  or  as 

long  as  wide,  almost  symmetrical,  thick,  flattened,  bluntly  conical  (fig.  27, 
h).  Anal  cirri  short,  thick,  almost  spherical.  Second  segment  with  seti- 
gerous lobe  and  setae  well  developed E.  longa 

Dorsal  cirri  much  larger  than  parapodial  lobes,  v/ider  than  long 3 

3.  Dorsal  cirri  slightly  asymmetrical,  oval  (fig.  27,  g).     Color  in  alcohol:  Dorsum 

brownish  with  bluish-violet  irridescence.  Anal  cirri  short,  thick,  almost 
spherical.     Second  segment  with  setigerous  lobe  and  setae  well  developed. 

E.  flava 
Dorsal  cirri  much  wider  than  long  (about  2.5  times),  asymmetrical,  obliquely 
oval,  with  lower  part  larger  than  upper  part  (fig.  27,  /).  Color  in  alcohol: 
Middle  third  of  dorsum  light  brownish,  lateral  third  deep  reddish  brown. 
Anal  cirri  elongate,  cylindrical,  3-4  times  longer  than  wide.  Second  segment 
with  setigerous  lobe  and  setae  poorly  developed  or  lacking. 

E.  spetsbergensis 


Subgenus  Mysta  Malmgren,  1865 
Eteone  {Mysta)  barbata  (IVIalmgren,  1865) 

Figure  27,  e 

Mysta  barbata  Malmgren,  1865,  p.  101,  pi.  15,  fig.  34;  1867,  p.  26,  pi.  3,  fig.  20.— 
Bergstrom,  1914,  p.  207,  fig.  79.— Southern,  1914,  p.  75.— Eliason,  1920, 
p.  28.— Augener,  1928,  p.  711;  1939,  p.  136.— Zatsepin,  1948,  p.  113,  pi.  29, 
fig.  14. — Wesenberg-Lund,  1951,  p.  31. 

Eteone  striata  Ditlevsen,  1917,  p.  66,  pi.  5,  figs.  11,  17,  19. 

Eteone  (Mysta)  barbata  Fauvel,  1923,  p.  176. — Annenkova,  1937,  p.  159;  1938, 
p.  146.— Berkeley  and  Berkeley,  1942,  p.  190.— Thorson,  1946,  p.  62,  fig. 
27,  A-B. 

Description. — Length  42-180  mm.;  width  without  setae  2.5-7  mm. 
See  key.  Color:  In  life:  Three  iridescent,  brownish-purple  dorsal 
longitudinal  bands,  median  and  lateral. 

New  records. — Arctic  Alaska:  Eluitkak  Pass,  Elson  Lagoon  near 
Point  Barrow,  stony-mud;  off  Point  Barrow  base,  6.1-27  fms.,  up 
to  3.2  miles  from  shore,  on  bottoms  of  mud,  gravel,  stones,  rocks; 
also  caught  in  trap  on  bottom  (3  stations,  4  specimens). 


234  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  los 

Distribution. — Scattered  records  in  the  Arctic :  Siberian  and  Alaskan 
Arctic,  Spitsbergen,  Novaya  Zemlya,  Kara  Sea.  Also  Iceland,  Nor- 
way to  Ireland,  North  Sea,  Baltic;  Bering  Sea;  north  Japan  Sea. 
In  5.5-60  fathoms. 


Eteone  longa  (Fabricius,  1780) 

FiGUKE  27,  h 

Nereis  longa  Fabricius,  1780,  p.  300. 

Eteone  longa  Oersted,  1843,  p.  185,  pi.  2,  figs.  20,  28. — Bergstrom,  1914,  p.  192, 
fig.  72.— Chamberlin,  1920,  p.  11.— Eliason,  1920,  p.  26.— Fauvel,  1923, 
p.  172,  fig.  62,  a-d;  1933,  p.  16.— Augener,  1928,  p.  710;  1939,  p.  136.— 
Annenkova,  1934,  p.  322;  1937,  p.  158;  1938,  p.  145.— Thorson,  1946,  p. 
59,  fig.  26,  A-E.— Gorbunov,  1946,  p.  38.— Zatsepin,  1948,  p.  113,  pi.  29, 
fig.  13. — Berkeley  and  Berkeley,  1948,  p.  41,  figs.  57,  58. — Wesenberg-Lund, 
1950a,  p.  11,  pi.  2,  figs.  6,  7;  1950b,  p.  38;  1951,  p.  29. 

Eteone  arctica  Malmgren,  1867,  p.  27,  pi.  2,  fig.  12. 

Eteone  cinerea  Webster  and  Benedict,  1884,  p.  705,  pi.  1,  figs.  1-5. 

Eteone  tuberculata  Treadwell,  1922,  p.  174,  figs.  7-10. 

Eteone  californica  Hartman,  1936a,  p.  131,  figs.  43-46;  1948,  p.  20,  fig.  4,  a-d. — 
Rioja,  1941,  p.  687. 

Description. — Length  10-65  mm.,  width  without  setae  0.5-3  mm. 
(length  up  to  160  mm. — Berkeley  and  Berkeley,  1948).  See  key. 
Color:  In  life:  White. 

Remarks. — E.  tuberculata  Treadwell  (1922)  from  Friday  Harbor, 
Washington,  E.  calijornica  Hartman  (1936)  from  central  California 
(type  examined  as  well  as  specimens  from  southeastern  Alaska — 
Hartman,  1948),  and  E.  cinerea  Webster  and  Benedict  (1884)  from 
Provincetown,  Massachusetts  (type  in  USNM),  appear  to  be 
synonymous  with  E.  longa. 

New  records. — Arctic  Alaska:  Eluitkak  Pass,  Elson  Lagoon  near 
Point  Barrow,  6.6  fms.;  off  Point  Barrow  base,  1.7-75.5  fms.,  up  to  8 
miles  from  shore,  on  bottoms  of  sandy-mud,  stones,  and  various 
combinations  of  mud,  pebbles,  stones,  gravel,  rocks  (12  stations,  40 
specimens).  West  Coast  North  America:  San  Juan  Archipelago, 
Washington,  Pettibone.  West  Greenland:  Vaigat,  Disko  Island, 
mud,  Bartlett,  1937. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian,  Alaskan, 
and  Canadian  Arctic,  Greenland,  Spitsbergea,  Franz  Josef  Land, 
Novaya  Zemlya.  Also  Iceland,  Norway  to  English  Channel; 
Hudson  Bay  to  Massachusetts;  Bering  Sea  to  Mexico;  north  Japan 
Sea;  China.     In  low  water  to  518  fathoms. 


]VIARINE    POLYCHAETE    WORMS — PETTIBONE  235 

Eteoneflava  (Fabricius,  1780) 

Figure  27,  g 

Nereis  flava  Fabricius,  1780,  p.  299. 

Eieone  flava  Malmgren,  1865,  p.  102,  pi.  15,  fig.  35.— Bergstrom,  1914,  p.  196, 

fig.  74.— Eliason,  1920,  p.  27.— Fauvel,  1923,  p.  173,  fig.  62,  e,  f.— Augener, 

1928,  p.  709.— Annenkova,  1937,  p.  158;  1938,  p.  145.— Zatsepin,  1948,  p. 

113,  pi.  29,  fig.  12.— Wesenberg-Lund,  1950a,  p.  11,  pi.  2,  fig.  7,  c;  1950b,  p. 

39;  1951,  p.  30. 
Eteone  depressa  Malmgren,  1865,  p.  103,  pi.  15,  fig.  36. — Southern,  1914,  p.  79. 
Eteone  sarsi  Malmgren,  1867,  p.  28,  pi.  2,  fig.  14. — Webster  and  Benedict,  1887, 

p.  711. 
Eteone  lentigera  Malmgren,  1867,  p.  29,  pi.  2,  fig.  13. 

Description. — Length  24-68  mm.;  width  without  setae  1-4  mm. 
(length  up  to  120  mm. — ^Fauvel,  1923).  See  key.  Color:  In  life: 
Pinkish  flesh  and  grayish  white. 

New  records. — ^Arctic  Alaska:  Eluitkak  Pass,  Elson  Lagoon  near 
Point  Barrow,  6.6  fms.;  off  Point  Barrow  base,  8.3-54.6  fms.,  up  to 
7.5  miles  from  shore,  on  bottoms  of  mud,  and  various  combinations 
of  mud,  stones,  gravel,  rock,  large  perforated  rocks,  shells  (10  stations, 
22  specimens). 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian  and 
Alaskan  Arctic,  Greenland,  Spitsbergen,  Novaya  Zemlya.  Also 
Iceland,  Faroes,  Norway  to  English  Channel;  east  coast  North  Amer- 
ica (Maine);  Bering  Sea;  north  Japan  Sea.  In  low  water  to  471 
fathoms. 

Eteone  spetsbergensis  Malmgren,  1865 

Figure  27,  / 

Eteone  spetsbergensis  Malmgren,  1865,  p.  102,  pi.  15,  fig.  38.— Th6el,  1879,  p.  31, 
pi.  2,  figs.  21,  22.— Bergstrom,  1914,  p.  202,  fig.  77.— Ditlevsen,  1917,  p.  66, 
pi.  5,  figs.  12,  14,  18.— Eliason,  1920,  p.  27.— Augener,  1928,  p.  708.— Annen- 
kova, 1937,  p.  159;  1938,  p.  145.— Hartmau,  1948,  p.  20,  fig.  5,  b.— Berkeley 
and  Berkeley,  1948,  p.  42.— Zatsepin,  1948,  p.  113,  pi.  29,  fig.  11.— Wesen- 
berg-Lund, 1951,  p.  31. 

Description. — Length  17-75  mm.,  width  without  setae  0.7-4  mm. 
(length  up  to  100  mm. — Augener,  1928).  See  key.  Color:  In  life: 
Pale  green  ventraUy,  reddish  brown  dorsolateral  bands,  with  dark 
dorsal  cirri;  orange  colored  eggs. 

New  records. — Arctic  Alaska:  Off  Browerville  near  Point  Barrow; 
off  Point  Barrow  base,  1.7-36  fms.,  up  to  A.)i  miles  from  shore,  on 
bottoms  of  sandy-mud  and  various  combinations  of  mud,  gravel, 
rocks,  stones,  shell  (4  stations,  4  specimens). 

Distribution. — Scattered  records  in  the  Arctic:  Arctic  Alaska,  Spits- 
bergen, Franz  Josef  Land,  Novaya  Zemlya.  Also  Iceland,  Norway 
to  Scotland;  Bering  Sea;  western  Canada;  north  Japan  Sea.  In  1.7- 
40  fathoms. 

261112—54 3 


236  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  103 

Eteone  sp.  larvae 

Larvae  of  a  species  of  Eteone  found  in  a  mass  of  jelly  (off  Point 
Barrow  base,  1.7  fms.,  300  ft,  out,  on  sandy-mud,  July  20,  1948);  11 
segments,  of  which  8  were  setigerous;  triangular  prostomium  with 
short  antennae  and  eyes;  tentacular  segment  with  short  tentacular 
cirri;  anal  cirri  short,  spherical. 


Genus  Phyllodoce  Savigny,  1817 

Subgenus  Anaitides  Czerniavsky,  1882 
Phyllodoce  (Anaitides)  groenlandica  Oersted,  1842 

Figure  27,  d,  i 

Phyllodoce  groenlandica  Oersted,  1842,  p.  121  {fide  Bergstrom,  1914);  1843,  p.  192, 
pi.  2,  figs.  19,  21,  22,  29-32.— Malmgren,  1865,  p.  96;  1867,  p.  21,  pi.  2,  fig. 
9.— Webster  and  Benedict,  1884,  p.  703;  1887,  p.  710.— Murdoch,  1885,  p. 
153.— Moore,  1903,  p.  428.— Mcintosh,  1908,  p.  86,  figs.— Ditlevsen,  1917, 
p.  56.— Fauvel,  1923,  p.  153,  fig.  54,  f-i.— Augener,  1928,  p.  703.— not 
Treadwell,  1937,  p.  28  {  —  Paranaitis  wahlbergi  (Malmgren),  in  USNM). — 
Annenkova,  1937,  p.  156.— Friedrich,  1939,  p.  122.— Thorson,  1946,  p.  52, 
fig.  21.— Gorbunov,  1946,  p.  38.— Berkeley  and  Berkeley,  1948,  p.  46,  fig. 
66.— Wesenberg-Lund,  1950a,  p.  10;  1950b,  p.  32;  1951,  p.  26. 

Phyllodoce  citrina  Moore,  1908,  p.  328. — Hartman,  1948,  p.  19  (part;  includes 
P.  groenlandica  and  P.  mucosa,  in  USNM;  not  P.  citrina  Malmgren,  1865). 

Anaitides  groenlandica  Bergstrom,  1914,  p.  141,  fig.  42. — Southern,  1914,  p.  68. — 
EUason,  1920,  p.  24  (part) .— Chamberiin,  1920,  p.  11.— Hartman,  1944a, 
pp.  335,  338;  1948,  p.  19.— Zatsepin,  1948,  p.  Ill,  pi.  29,  fig.  2. 

Description. — Length  16-285  mm.,  width  without  setae  1-9  mm. 
(length  up  to  450  mm. — Ditlevsen,  1917).  Segments  very  numerous. 
Body  elongate,  large,  robust,  linear,  flattened,  attenuate  posteriorly. 
Prostomium  cordiform,  notched  posteriorly  with  an  occipital  tubercle 
in  the  notch,  with  foiu-  short  subulate  frontal  antennae,  with  smaU, 
paired,  lateral,  retractile,  nuchal  knobs  just  anterior  to  the  first  pair 
of  tentacular  cirri.  Tentacular  cirri  unequal;  two  pairs  short,  two 
pairs  longer,  extending  to  segments  9-11.  Dorsal  cirri  of  median 
region  up  to  two  times  longer  than  wide,  subquadrangular,  with  distal 
ends  truncate;  ventral  cirri  oval,  with  asymmetrical  acuminate  tip. 
Anterior  part  of  extended  proboscis  transversely  rugose,  crowned 
mth  17  papillae;  basal  part  with  12  longitudinal  rows  of  smaU,  oval 
papillae  (6  rows  on  each  side),  with  10-20  papillae  per  row.  Color: 
In  life  and  in  alcohol:  Body  with  bluish  iridescence,  usually  heavily 
pigmented  dark  greenish  brown  or  deep  reddish  blue,  with  dorsal  and 
ventral  cirri  tan  to  dark  brown;  on  some,  body  not  so  darkly  pig- 
mented, with  cirri  only  partly  pigmented. 


MARINE    POLYCHAETE    WORMS — PETTIBONE 


237 


Figure  27. — Phyllodocidae:  a,  Eteone,  dorsal  view  anterior  end;  b,  My  slides  borealts, 
dorsal  view  anterior  end;  c,  Eumida  minuta,  dorsal  view  anterior  end;  d,  Phyllodoce 
groenlandica,  dorsal  view  anterior  end;  e,  Eteone  barhata,  parapodium;  /,  Eteone  spets- 
bergensis,  parapodium;  g,  Eteone  flava,  parapodium;  h,  Eteone  longa,  parapodium;  i, 
Phyllodoce  groenlandica,  parapodium.     (For  explanation  of  symbols,  see  p.  210.) 


238  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  103 

Remarks. — Some  smaller  specimens  were  at  first  placed  under  P. 
citrina  Malmgren,  since  the  papillae  on  the  proboscis  were  fewer  than 
normal,  four  rows  on  each  side,  with  3-5  papillae  per  row.  An  inter- 
mediate-sized specimen  from  Canoe  Bay,  Alaska  (Hartman,  1948,  as 
P.  citrina)  has  five  rows  on  each  side,  with  as  many  as  6  papillae  per 
row.  It  appears  that  the  papillae  are  added  on  gradually  and  that 
only  in  the  fully  developed  individual  is  the  full  complement  of  6 
rows  of  papillae  on  each  side  developed ;  the  number  per  row  is  varia- 
ble, ranging  from  10  (^vith  some  spaces  as  if  some  had  dropped  off) 
to  20. 

New  records. — Arctic  Alaska:  Eluitkak  Pass,  Elson  Lagoon  near 
Point  Barrow;  Point  Barrow  base,  washed  ashore;  off  Point  Barrow 
base,  up  to  12.1  miles  from  shore,  1.7-123.5  fms.,  on  bottoms  of  mud, 
sandy-mud,  mass  of  worm  tubes,  and  various  combinations  of  mud, 
gravel,  stones,  pebbles,  rocks,  large  perforated  rocks,  shells;  in  screen 
trap  on  bottom  (23  stations,  74  specimens).  West  Greenland: 
Vaigat,  Disko  Island,  mud,  Bartlett,  1937.  East  Coast  North 
America:  Off  Labrador,  70  fms.,  Blue  Dolphin  Expedition,  1950;  off 
Maine,  Massachusetts,  intertidal  to  42  fms.,  U.  S.  Fish  Commission. 
West  Coast  North  America:  Washington  Sound,  Pettibone. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian,  Alaskan, 
and  Canadian  Arctic,  Davis  Strait,  Greenland,  Spitsbergen,  Franz 
Josef  Land,  Barents  Sea,  Novaya  Zemlya,  Kara  Sea;  Also  Iceland, 
Faroes,  Norway  to  English  Channel;  Hudson  Bay  to  Massachusetts; 
Bering  Sea  to  Washington;  north  Japan  Sea.  In  low  water  to  800 
fathoms. 

Genus  Eumida  Malmgren,  1865 
Eumida  minuta  (Ditlevsen,  1917) 

Figure  27,  c 

Eulalia  minuta  Ditlevsen,  1917,  p.  56,  pi.  4,  figs.  10,  12,  14. —  ?  Gorbunov,  1946, 

p.  38. 
Eulalia  arctica  Aunenkova,  1946,  pp.  185,  187,  fig.  1,  a-c. 

Description. — Length  1-8  mm.;  width  without  setae  0.3-0.8  mm. 
Segments  few  (12-36).  Body  very  small,  slightly  tapered  anteriorly 
and  posteriorly,  flattened  dorsoventrally.  Prostomium  semiglobular 
to  bluntly  conical,  with  two  large  eyes  near  posterior  border,  with 
antennae  rather  long,  slender,  filamentous,  almost  as  long  as  pro- 
stomium. First  tentacular  achaetous  segment  not  distinct  dorsally, 
with  first  pair  tentacular  cirri  lateral  to  prostomium  at  same  level  as 
the  eyes  (thus  placed  under  Eumida  and  not  Eulalia  Savigny);  two 
pairs  tentacular  cirri  on  second  segment  (first  setigerous);  fourth 
tentacular  cirri  on  third  segment;  tentacular  cirri  enlarged  basally, 


MARINE    POLYCHAETE    WORMS — PETTIBONE  239 

with  slender  filamentous  tips,  the  latter  about  as  long  as  the  enlarged 
basal  part  except  for  the  ventral  tentacular  cirri  on  the  second  segment 
which  are  much  shorter.  Dorsal  cirri  oval  or  bluntly  conical. 
Ventral  cirri  elongate  oval.  Anal  cirri  short,  oval,  1.5  to  2  times 
longer  than  wide.  Color:  In  alcohol:  Greenish  tan  to  brown.  In 
life:  Flesh  ventrally,  green  and  flesh  dorsally,  with  salmon-colored 
eggs  showing  through. 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  16.7-75.5 
fms.,  up  to  8  miles  from  shore,  on  various  combinations  of  gravel, 
small  stones,  rocks,  bryozoans,  hydroids,  Saxicava  shells  (6  stations, 
46  specimens). 

Distribution. — Few  scattered  records  in  the  Arctic:  Siberian  and 
Alaskan  Arctic,  Davis  Strait.     In  16.7-75.5  fathoms. 

Family  Hesionidae 

Prostomium  usually  with  four  eyes,  two  or  three  antennae,  two 
biarticulate  palps  (fig.  28,  a).  Two  to  eight  pairs  tentacular  cirri  on 
one  to  four  more  or  less  distinct  achaetous  tentacular  segments. 
Parapodia  biramous  or  subbiramous;  notopodia  may  be  greatly  re- 
duced, with  notosetae  simple  or  lacking;  neurosetae  compound  (fig. 
28,  b).  Dorsal  cirri  long,  more  or  less  distinctly  articulated;  ventral 
cirri  shorter;  two  long  anal  cirri.  Proboscis  cylindrical,  eversible, 
with  marginal  papiUae,  with  or  without  horny  jaws. 

Represented  by  a  single  species  from  Point  Barrow.  (The  speci- 
mens from  Point  Barrow,  referred  to  Castalia  multipapillata  Theel  by 
Murdoch  (1885),  do  not  agree  with  this  species  nor  with  any  other 
species  of  hesionid;  the  parapodia  are  biramous,  both  lobes  being 
pointed,  the  notopodia  are  well  developed,  with  capillary  setae; 
neurosetae  compound,  with  delicate  tips;  the  proboscis  has  numerous 
small  papillae;  tentacular  cirri  four  pairs;  prostomium  with  two 
pairs  eyes,  pair  biarticulate  palps,  antennae  ?  (missing  or  absent); 
the  specimens  are  very  small  and  in  poor  condition.) 

Genus  Castalia  Savigny,  1820;  emend.  Fauvel,  1923 
Castalia  aphroditoides  (Fabricius,  1780) 

Figure  28,  a,  h 

Nereis  aphroditoides  Fabricius,  1780,  p.  296. 

Castalia  ardica  Malmgren,  1867,  p.  32. — Annenkova,  1931,  p.  203. 

Castalia  fabricii  Malmgren,  1867,  p.  32.— Th6el,  1879,  p.  37,  pi.  3,  figs.  36,  37.— 

Berkeley  and  Berkeley,  1943,  p.  130. 
Castalia  aphroditoides  Wir6n,  1883,  p.  401.— Augener,  1913,  p.  260;  1928,  p.  715.— 

Zatsepin,  1948,  p.  114.— Wesenberg-Lund,  1950a,  p.  13,  pi.  3,  fig.  14;  19o0b, 

p.  44. 
Psammate  aphroditoides  Chamberlin,  1920,  p.  13. 


240  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  103 

Description. — Length  2.5-18  mm.,  width  without  setae  0.7-2.5  mm. 
Segments  25-43.  Body  cylindrical.  Prostomium  rectangular,  wider 
than  long,  widest  anteriorly,  notched  posteriorly,  with  two  pairs 
eyes  in  trapezoidal  arrangement,  anterior  pair  larger;  palps  on  an- 
terolateral borders  of  prostomium;  two  filiform  antennae  medial  to 
palps  (without  unpaired  antenna);  antennae  subequal,  about  same 
length  as  prostomium.  SLx  pairs  tentacular  cirri  on  three  more  or  less 
distinct  achaetous  segments;  tentacular  cirri  long,  filiform,  articulated, 
with  prominent  cirrophores.  Without  distinct  notopodia  (acicula  in 
cirrophores  of  dorsal  cirri  only);  without  notosetae.  Neuropodia 
well  developed,  with  three  diverging  conical  distal  lobes :  Lip  anterior 
to  setae  with  median  and  dorsal  conical  lobes;  lip  posterior  to  setae 
with  rounded  median  and  conical  ventral  lobes;  neurosetae  compound. 
Dorsal  and  anal  cirri  long,  articulated.  Proboscis  barrel-shaped  with 
10  soft  papillae  around  opening  (four  dorsal  to  lateral  grooves,  three 
each  on  two  ventrolateral  folds),  with  a  median  ventral  notch, with 
two  thickened  ventrolateral  ridges  (the  so-called  jaws)  lateral  to  the 
ventral  notch.  Color:  In  alcohol:  Without  color,  or  slightly  greenish 
dorsally  (especially  in  anterior  part),  or  brownish.  In  life:  Orange- 
flesh,  with  greenish  eggs  inside,  with  red  eyes. 

Remarks. — Two  young  specimens  found  in  plankton  tow  beneath 
ice  February  28,  1950. 

New  records. — Arctic  Alaska:  Eluitkak  Pass,  Elson  Lagoon  near 
Point  Barrow,  6.6  fms.,  stony-mud;  off  Point  Barrow  base,  21-75.5 
fms.,  up  to  8  miles  from  shore,  on  bottoms  of  mud,  stones,  and  various 
combinations  of  mud,  stones,  gravel,  rocks,  and  from  breaking  apart 
fohaceous  brj^ozoans  (10  stations,  28  specimens).  Northwest 
Greenland:  Walrus  grounds,  Murchison  Sound,  Bartlett,  1938. 
East  Coast  North  America:  Off  Labrador,  45  fms..  Blue  Dolphin 
Expedition,  1949. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian,  Alaskan, 
and  Canadian  Arctic,  Greenland,  Spitsbergen,  Franz  Josef  Land, 
Novaya  Zemlya,  Kara  Sea.  Also  Bering  Sea;  Denmark;  Hudson  Bay, 
Labrador.     In  2-75.5  fathoms. 

Family  Syllidae 

Body  of  small  size.  Prostomium  with  four  eyes  (sometimes  six), 
two  palps  (may  be  reduced  and  fused),  three  antennae  (figs.  28,  29). 
Tentacular  segment  achaetous,  with  one  or  two  pairs  tentacular  cirri. 
Parapodia  uniramous,  with  dorsal  and  ventral  cirri  (latter  may  be 
absent).  Two  anal  cirri.  Setae  mostly  compound,  sometimes  with 
additional  simple  setae.     Proboscis  eversible,  armed  or  not  with  one 


MARINE    POLYCHAETE    WORMS — PETTIBONE  241 

to  several  chitinous  teeth.  Reproduction  direct  (epigamy)  or  by 
stolons,  sometimes  having  sexual  dimorphism;  with  swimming  capillary 
setae  at  maturity. 

Represented  by  6  genera  and  11  species. 


Key  to  the  genera  of  Syllidae  from  Point  Barrow 

1.  Without  ventral  cirri.     Palps  reduced,  united,  turned  under  on  ventral  side 

of  prostomium  (except  for  forked  palps  of  male  stolon;  fig.  29,  a).  Antennae 
and  dorsal  cirri  smooth  or  faintly  annulated.  Reproduction  by  stolons; 
stem  forms  produce  by  transverse  fission,   singly  or  in   chains,   sexually 

dimorphic  male  and  female  forms  with  swimming  setae Autolytus — 2 

With  ventral  cirri  (fig.  28,  d) .     Palps  better  developed,  free  or  fused 4 

2.  Prostomium  normal,  with  4  dorsal  eyes  in  trapezoidal  arrangement,  with  3 

antennae,  short  fused  palps;  2  pairs  tentacular  cirri  (fig.  29,  a).  Without 
swimming  setae.     Pharynx  long,  more  or  less  sinuous,  usually  with  a  crown 

of  teeth Stem  form  or  stock  of  Autolytus  (p.  242) 

Prostomium  abnormal,  with  4  large  eyes  (usually  2  dorsolateral  and  2  ventro- 
lateral) ;  2-3  pairs  tentacular  cirri  (fig.  29,  c-f) .  Body  divided  into  an  anterior 
unmodified  region,  a  middle  region  with  long  swimming  setae  and  well- 
developed,  paddlelike  parapodia,  and  with  or  without  a  posterior  unmodified 
region.     Without  pharynx Sexual  stolons  of  Autolytus — 3 

3.  Prostomium  with  3  antennae  (very  long  median  one  and  pair  of  small  frontal 

ones)  and  pair  of  anterior  bifurcate  palps;  1  pair  tentacular  cirri  very  long, 
similar  to  median  antennae  (these  may  be  the  dorsal  cirri  of  the  first  setiger 
or  of  a  small,  knoblike  achaetous  lobe;  fig.  29,  e,f).  Testes  and  sperm  con- 
fined to  anterior  unmodified  region. 

Male  form  of   Autolytus    (Polybostrichus)  (p.  242) 
Prostomium  with  3  subequal  antennae,  without  palps;  none  of  tentacular 
cirri  especially  elongate  (fig.  29,  c,  d).     Most  of  body  filled  with  eggs;  eggs 
carried  in  large  sac  on  ventral  surface  (gestation). 

Female  form  of  Autolytus  (Sacconereis)  (p.  242) 

4.  Dorsal  cirri  distinctly  moniliform  throughout  body  (fig.  28,  c) .     Palps  separate 

for  their  entire  length  or  fused  basally.  Reproduction  direct  or  by  single 
stolons.  Sexual  stolons  (Chaetosyllis) :  Small  prostomium,  with  2  pairs 
eyes,  pair  short  moniliform  antennae;  without  tentacular  cirri;  1  anterior 
unmodified  segment,  a  middle  region  with  long  swimming  setae,  and  few 

unmodified  posterior  segments Syllis  (p.  252) 

Dorsal  cirri  smooth  or  indistinctly  articulate  (may  be  more  or  less  articulate 
anteriorly) .     Palps  fused  at  base  only  or  throughout .__  5 

5.  Palps  fused  for  nearly  their  entire  length  (fig.  28,  k-m).     Body  very  small. 

Antennae  and  dorsal  cirri  short,  not  moniliform.  One  pair  tentacular  cirri 
(may  be  rudimentary).  Reproduction  generally  direct,  with  swimming 
setae  at  maturity;  eggs  and  larvae  fixed  to  dorsal  or  ventral  surface  of 

female 6 

Palps  fused  at  base  only  (fig.  28,  g).  Body  larger.  Antennae  and  dorsal 
cirri  longer,  smooth  (antennae  and  anterior  dorsal  cirri  may  be  indistinctly 
articulate,  especially  distally).  Two  pairs  tentacular  cirri.  Reproduction 
direct,  with  swimming  setae  at  maturity 7 


242  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  103 

6.  Antennae  and  dorsal  cirri  swollen  at  base,  tapering  to  narrow  tip  (fig.  28,  m) . 

Body  and  parapodia  usually  covered  with  adhesive  papillae  and  fine,  granular 

material Sphaerosyllis  (p.  255) 

Antennae  and  dorsal  cirri  clavate  or  conical  (fig.  28,  k,  I).  Bodj'  and  parapodia 
not  covered  with  papillae Exogone  (p.  257) 

7.  Compound  neurosetae  with  distal  blades  all  rather  short  (fig.  28,  i).     Pro- 

boscis with  a  distal  and  proximal  row  of  soft  papillae,  with  distal  circular 

chitinous  margin  denticled  (fig.  28,  g,  h) Eusyllis  (p.  259) 

Compound  neurosetae  with  some  distal  blades  elongate  (fig.  28,  j).  Proboscis 
with  distal  row  of  soft  papillae  only,  with  distal  circular  chitinous  margin 
smooth Pionosyllis  (p.  262) 


Genus  Aiitolytus  Grube,  1850 

In  working  over  the  rather  numerous  specimens  of  Autolytus  in 
the  collections  from  Point  Barrow,  several  things  were  revealed  which 
throw  light  on  the  confusion  in  this  group  caused  by  the  formation  of 
sexually  dimorphic  stolons.  For  correct  description  of  any  species, 
one  should  have  the  stem  form,  the  sexual  buds  in  the  process  of 
formation  including  the  mature  sexual  buds  attached  to  the  stem 
form,  and  the  mature  male  and  female  stolons  separated  from  the 
stem.  Very  often,  however,  stem  forms  and  the  male  and  female 
stolons  of  one  species  have  been  described  and  given  different  names, 
while  sexual  stolons  and  stem  forms  of  different  species  have  been 
given  the  same  name.  A.  fallax  Malmgren,  described  originally  from 
specimens  from  Spitsbergen,  is  of  particular  interest.  Malmgren 
described  and  figured  the  stem  form  -with  the  head  of  a  male  stolon 
forming  between  se tigers  13  and  14  and  indicated  that  the  sexual 
stolons  were  not  known.  A.  prismaticus  (Fabricius),  described 
originally  from  specimens  from  Greenland,  has  a  characteristic  color 
pattern  of  three  dark  longitudinal  bands  (median  and  lateral  at  the 
level  of  the  bases  of  the  dorsal  cirri).  The  sexual  stolons  have  six 
unmodified  setigers  anterior^toHhe  setigers' with  swimming  setae;  the 
stolons  show  the  characteristic  color  pattern  of  the  stem  form.  In 
the  absence  of  color,  it  would  be  difficult  to  distinguish  A.  prismaticus 
from  certain  other  species  of  Autolytus.  It  apparently  has  been  the 
usual  practice  to  identify  any  sexual  stolon  of  unknown  connections 
that  has  six  anterior  unmodified  setigers  as  A.  prismaticus,  and  some- 
times to  identify  the  male  stolons  as  Polyhostrichus  longosetosus 
Oersted  or  A.  longisetosus  (Oersted).  However,  as  found  in  the  Point 
Barrow  material,  A.  fallax  is  the  most  common  species  of  Autolytus 
and  it  was  found  along  \vith  A.  prismaticus  and  A.  alexandri.  The 
majority  of  the  specimens  of  A.  Jallax  were  in  the  process  of  stolon 
formation,  the  head  of  the  stolon  being  formed  between  setigers  13 


MARINE    POLYCHAETE    WORMS — PETTIBONE 


243 


Figure  28. — Hesionidae:  a,  Castalia  aphroditoides,  dorsal  view  anterior  end;  b,  same, 
parapodium,  posterior  view.  Syllidae:  c,  Syllis  fasciata,  dorsal  view  anterior  end;  d, 
same,  parapodium;  e,  same,  neuroseta;  /,  Syllis  cornuia,  neurosetae,  with  short  (1)  and 
long  (2)  blades;  g,  Eusyllis  blomstrandi,  dorsal  view  anterior  end,  proboscis  partially 
extended;  h,  same,  frontal  view  extended  proboscis  showing  distal  row  of  papillae  and 
denticled  margin;  t,  same,  neuroseta;  /,  Pionosyllis  compacta,  neurosetae,  lower  one  (1) 
with  shorter  blade,  upper  one  (2)  with  longer  blade;  k,  Exogoru  dispar,  dorsal  view  anterior 
end;  /,  Exogone  naidina,  dorsal  view  anterior  end;  m,  Sphaerosyllis  erinaceus,  dorsal  view 
anterior  end.     (For  explanation  of  symbols,  see  p.  210.) 


244  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  103 

and  14.  Although  no  mature  sexual  buds  were  found  attached  to  the 
stem  form,  the  rudiments  of  the  notopodia  with  the  developing  swim- 
ming setae  could  be  seen,  beginning  on  setiger  7.  Male  and  female 
stolons  with  six  anterior  unmodified  segments  were  found.  Of  these, 
two  female  stolons  showed  the  characteristic  banding  of  A.  prismaticus 
(this  banding  was  noted  on  aU  the  developing  sexual  buds  still  attached 
to  the  stem  forms  of  this  species).  The  others  did  not  show  this 
banding  and  are  considered  to  be  the  sexual  stolons  of  A.  jallax.  A. 
Jallax,  then,  along  with  A.  prismaticus,  has  six  anterior,  unmodified 
setigers  and  manj^  of  the  records  of  A.  prismaticus  in  the  literature 
will  no  doubt  prove  to  be  A.  jallax. 

The  stem  form  of  A.  fallax  has  been  referred  to  A.  prolijer  (Miiller) 
by  Augener  (1913,  1928)  and  Wesenberg-Lund  (1947),  thus  the  stem 
form  of  A.  fallax  has  been  confused  wdth  A.  prolifer  and  sexual  forms 
with  A.  prismaticus. 

There  appears  to  be  a  correlation  in  the  place  of  formation  of  the 
sexual  stolons  in  the  stem  form  with  the  number  of  anterior,  unmodi- 
fied setigers  and  the  number  of  tentacular  cirri  in  the  sexual  stolons. 
At  least  for  a  group  of  species,  including  A.  prismaticus,  A.  fallax,  A. 
cornutus  A.  Agassiz,  and  A.  ornatus  (Verrill,  1873),  the  stem  forms 
produce  sexual  stolons  singly,  with  the  head  of  the  stolon  forming 
between  setigers  13  and  14.  The  stolon,  then,  consists  of  the  segments 
of  the  stem  posterior  to  setiger  13  (differing  from  A.  prolifer  where 
the  sexual  stolons  are  proliferated  more  posteriorly,  often  in  chains  of 
2  to  8).  The  posterior  part  develops  gradually  into  the  male  or 
female  sexual  stolon  and  breaks  off  when  mature.  The  anterior  13 
setigers  of  the  stem  regenerates  another  posterior  end.  The  sexual 
stolons  of  these  four  species  have  six  mimodified  setigers  anterior  to 
the  swimming  setae  and  usually  a  more  or  less  developed  immodified 
posterior  region.  The  stolons  have  three  pairs  of  tentacular  cirri  on 
the  first  two  achaetous  tentacular  segments;  the  first  two  pairs  on  the 
first  segment  and  the  third  pair  on  the  second  segment.  The  upper 
pair  of  tentacular  cirri  on  the  first  segment  are  similar  to  the  dorsal 
cirri.  The  lower  pair  may  be  very  short,  about  the  length  of  the 
prostomium.  The  third  pair  of  tentacular  cirri  in  the  female  are 
similar  to  the  dorsal  cirri;  in  the  male  they  are  very  long,  similar  to 
the  median  antenna.  In  addition,  there  are  a  pair  of  small  achaetous 
lobes.  In  contrast,  in  A.  prolifer  the  sexual  stolons  have  three  ante- 
rior unmodified  setigers  (sometimes  two  or  four)  and  a  poorly  devel- 
oped posterior  region;  there  is  a  single  tentacular  achaetous  segment 
with  two  pairs  of  tentacular  cirri  (in  the  female  both  the  upper  and 
lower  pairs  are  short;  in  the  male  the  lower  one  is  similar  to  the  dorsal 
cirri  and  the  upper  one  is  very  long,  similar  to  the  median  antenna). 
In  A.  alexandri  the  sexual  stolons  have  14  (sometimes  13)  anterior 


MARINE    POLYCHAETE    WORMS — PETTIBONE  245 

unmodified  setigers  and  a  well-developed  posterior  region;  there  is  a 
single  achaetous  tentacular  segment  with  two  pairs  of  tentacular  cirri; 
the  first  setigerous  segment  in  the  female  is  similar  to  the  following 
segment,  but  in  the  male  it  has  a  very  long  pair  of  dorsal  cirri  similar 
to  the  median  antenna.  Thus  the  male  stolons  of  the  different  species 
of  Autolytus  agree  in  having  a  very  long  pair  of  anterior  cirri  similar 
to  the  median  antenna,  but  they  may  be  developed  as  the  upper  pair 
of  tentacular  cirri  of  a  single  tentacular  segment  as  in  ^.  prolifer,  as 
the  dorsal  cirri  of  the  first  setigerous  segment  as  in  ^.  alexandri,  or  as 
the  third  pair  of  tentacular  cirri  on  the  second  tentacular  segment 
just  dorsal  to  a  pair  of  achaetous  lobes  as  in  A.  prismaticus. 

The  number  of  setigers  in  the  anterior,  unmodified  region  of  the 
sexual  stolons  appears  to  be  a  good  taxonomic  character  for  most 
species.  That  the  number  is  somewhat  variable  in  A.  prolifer  (usually 
three,  sometimes  two  and  four)  does  not  mean  that  it  is  of  no  signifi- 
cance in  other  forms.  Thus  the  specimens  of  Autolytus  with  six 
anterior  unmodified  setigers,  which  Dales  (1951)  referred  to  A.  prolifer, 
are  no  doubt  another  species.  The  description  agrees  well  with  that 
of  A.  cornutus. 

All  three  species  of  Autolytus  from  Point  Barrow  have  the  body 
linear  elongate,  attenuated  posteriorly,  flattened  ventrally,  arched 
dorsally.  Prostomium  with  four  eyes  in  trapezoidal  arrangement, 
with  palps  fused,  turned  ventrally,  exceeding  the  prostomium  only 
sUghtly.  Neuropodia  with  composite  setae  with  blades  short,  biden- 
tate,  and  usually  with  a  simple  bayonette  seta. 

Key  to  the  species  of  Autolytus  from  Point  Barrow 

1.  Stem  Form:  Dorsal  cirri  irregular  in  length,  at  least  some  longer  than  body 
width,  pigmented  with  reddish  globules;  body  whitish,  brownish,  or  trans- 
versely banded  with  reddish  globules,  2  narrow  bands  per  segment;  with 
nuchal  epaulettes  on  first  2  (to  4)  setigers;  sexual  buds  formed  singly  (?), 
the  heads  forming  posterior  to  setiger  14.  Sexual  Stolons:  14  setigers 
anterior  to  swimming  setae;  nuchal  epaulettes  on  first  3  setigers;  2  pairs 
tentacular  cirri,  the  upper  one  longer;  in  male,  first  pair  dorsal  cirri  on  first 
setiger  very  long,  similar  to  median  antenna;  female  stolons  with  2-lobed 

egg  sac A.  alexandri 

Stem  Form:  Dorsal  cirri,  except  anterior  2  pairs,  shorter  than  body  width, 
not  extending  beyond  or  only  slightly  be3'ond  the  setae,  subequal;  sexual 
buds  formed  singly,  the  heads  forming  between  setigers  13  and  14.  Sexual 
Stolons:  6  setigers  anterior  to  swimming  setae;  3  pairs  tentacular  cirri;  in 
male,  first  pair  similar  to  dorsal  cirri,  with  second  short  pair  at  its  ventral 
base,  third  pair  very  long  and  stout  (similar  to  median  antenna)  with  small 
achaetous  lobe  at  its  ventral  base  (fig.  29,  e,  f) ;  in  the  female,  similar  except 
both  the  first  and  third  pairs  are  similar  to  the  following  dorsal  cirri  (fig. 
29,  c,  d);  female  stolons  with  1-lobed  egg  sac 2 


246  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  los 

2.  Stem  Form:  Usually  with  2  dorsolateral  longitudinal  black  to  brown  bands 
just  dorsal  to  bases  of  dorsal  cirri  (may  be  faint  or  absent);  with  short 
semicircular  epaulettes  on  tentacular  segment  only.  Sexual  Stolons: 
Colorless  or  female  stolon  dusk}'^,  especially  posteriorly;  male  stolon  rusty 
brown,    darkest    on   anterior    region,    especially   laterally;   epaulettes    on 

tentacular  segment  only  (fig.  29,  c-f) A.  fallax 

Stem  Form:  With  3  longitudinal  black  bands,  1  medium  and  2  lateral  at  same 
level  as  bases  of  dorsal  cirri;  medium  band  continuous  throughout  length  of 
body,  lateral  ones  may  fade  out  in  anterior  fourth  or  be  lacking;  with  semi- 
circular epaulettes  on  tentacular  segment  and  part  of  first  setiger  (fig.  29,  a). 
Sexual  Stolons:  With  3  longitudinal  pigmented  bands,  median  one  con- 
tinuous throughout  most  of  body,  lateral  ones  along  bases  of  dorsal  cirri 
in  anterior  unmodified  region;  with  epaulettes  on  tentacular  segment  and 
part  of  first  setiger A.  prisxnaticus 

Autolytus  alexandri  Malmgren,  1867 

Autolytus  alexandri  Malmgren,  1867,  p.  37,  pi.  7,  fig.  39. — Verrill,  1881,  p.  292, 

pi.  12,  fig.  8.— Chamberlin,  1920,  p.  12.— Friedrich,  1939,  p.  123.— Hartman, 

1942a,  p.  13;  1944a,  p.  338,  pi.  13,  fig.  11  (not  pi.  13,  fig.  2);  1945,  p.  17, 

pi.  2,  fig.  11. 
Stephanosyllis  pida  Verrill,  1874a,  pp.  43,  132,  pi.  4,  fig.  6;  1874b,  pp.  361,  362, 

pi.  4,  fig.  1  (not  Proceraea  picta  Ehlers,  1864). 
Stephanosyllis  ornata  Verrill,  1874a,  p.  132  (not  Proceraea  ornata  Verrill,  1873). 
Proceraea    (Stephanosyllis)    ornata   Webster   and   Benedict,    1887,   p.    724    (not 

Proceraea  ornata  Verrill,  1873). 
Autolytus  verrilli  Marenzeller,  1892,  p.  416,  pi.  19,  fig.  4. — Augener,  1928,  p.  726. — 

Wesenberg-Lund,  1947,  p.  33,  figs.  14,  15;  1950a,  p.  18;  1950b,  p.  52;  1951, 

p.  52. 

Description. — Stem  form:  Length  4-12  mm.,  width  without  setae 
0.6-0.8  mm.,  up  to  35-64  segments.  Antennae  rather  short,  thick, 
subequal  or  median  one  up  to  twice  the  length  of  the  lateral;  tentac- 
ular cirri  equal  to  or  shorter  than  antennae,  upper  pair  longer; 
first  pair  dorsal  cirri  as  long  as  or  longer  than  median  antenna;  rest 
of  dorsal  cirri  rather  irregular,  unequal,  at  least  some  longer  than  body 
width,  with  cirrophores  prominent.  Distinct  epaulettes  on  first  two 
setigerous  segments  (or  to  end  of  setiger  4 — Hartman,  1945).  None 
showed  sexual  bud  formation  (sexual  bud  is  formed  between  segments 
25  and  26 — Hartman,  1945).  Color:  In  alcohol:  Colorless  or  trans- 
versely banded  with  reddish  to  brownish  granules,  may  be  two  bands 
per  segment. 

Female  stolon  (Sacconereis) :  A  single  specimen  of  72  setigers, 
16  mm.  long,  1.2  mm.  wide  without  setae,  3.5  mm.  wide  with  swimming 
setae,  consisting  of  14  anterior,  26  middle,  and  32  posterior  setigers. 
Prostomium  rounded,  with  four  large  dorsal  eyes,  three  antennae 
rather  short,  subequal;  two  pairs  tentacular  cirri  of  nearly  same 
length  as  antennae;  dorsal  cirri  of  anterior  part  of  unequal  length  but 
all  much  longer  than  body  width.  Thick  epaulettes  extending  on 
first  three  setigers.     The  stolon,  with  swimming  setae  and  fiUed  with 


MARINE    POLYCHAETE    WORMS — PETTIBONE  247 

eggs,  was  dredged  in  27  fathoms  February  18,  1950.  (A  2-lobed 
egg  mass  is  carried  ventrally — Wesenberg-Lund,  1947.) 

Male  stolon  (Polyhostrichus) :  No  specimens  taken  from  Point 
Barrow.  A  specimen  taken  in  plankton  at  Portage  Bay,  Alaska,  was 
12  mm.  long,  1.5  mm.  wide  without  setae,  4  mm.  wide  including 
swimming  setae,  with  61  setigers  consisting  of  14  anterior,  36  median, 
and  11  posterior  segments.  Prostomium  similar  to  A.  fallax;  two 
pairs  tentacular  cirri,  upper  pair  longer.  First  pair  dorsal  cirri  on 
first  setiger  very  long,  similar  to  median  antenna  (achaetous — ^Wesen- 
berg-Lund,  1947);  dorsal  cirri  in  anterior  region  long,  unequal;  in 
middle  region  short,  about  one-third  length  of  elongated  parapodial 
lobes.     Large  epaulettes  on  first  three  setigers. 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  16.7-123.5 
fms.,  up  to  15  mUes  from  shore,  on  bottoms  of  stones,  worm  tubes,  and 
various  combinations  of  mud,  pebbles,  gravel,  stones,  rocks,  large 
perforated  rocks,  with  bryozoans,  hydroids,  shells,  and  worm  tubes 
(17  stations,  53  specimens).  West  Coast  North  America:  Albatross 
surface  station,  Portage  Bay,  Alaska;  San  Juan  Channel,  Washington 
Sound,  25  fms.,  Pettibone,  1940.  East  Coast  North  America:  Off 
Labrador,  30-40  fms..  Blue  Dolphin  Expedition,  1949;  Woods  Hole 
region,  Massachusetts,  Pettibone,  1950. 

Distribution. — Scattered  records  in  the  Arctic:  Arctic  Alaska, 
Greenland,  Spitsbergen,  Franz  Josef  Land,  Barents  Sea.  Also  Iceland; 
Labrador  to  North  Carolina;  Bering  Sea  to  Washington.  In  low 
water  to  123.5  fathoms;  sexual  forms  at  surface. 

Autolytus  fallax  Malmgren,  1867 

Figure  29,  c-f 

?  Polyhostrichus  longosetosus  Oersted,  1843,  p.  183,  pi.  5,  figs.  62,  67,  71. — Tread- 

weU,  1937,  p.  28. 
Autolytus  fallax  Malmgren,  1867,  p.  33,  pi.  6,  fig.  41.— Ditlevsen,  1929,  p.  17. 
?  Autolytus  longisetosus  Malmgren,  1867,  p.  34,  pi.  7,  fig.  38. 
Autolytus  prolifer  Augener,  1913,  p.  258;  1928,  p.  724. — Wesenberg-Lund,  1947, 

p.  19,  figs.  8,  9  (part;  not  Nereis  prolifera  Miiller,  1788). 
Autolytus  prismaticus  Wesenberg-Lund,  1947,  p.  24  (part;  not  Nereis  prismatica 

Fabricius,  1780). 

Description. — Stem  form:  Length  3-18  mm.,  width  without  setae 
0.3-0.7  mm,,  up  to  45-78  segments.  Median  antenna  and  first  pair 
dorsal  cirri  very  long;  lateral  antennae  and  upper  tentacular  cirri 
about  half  as  loDg;  lower  pair  tentacular  cirri  and  second  pair  dorsal 
cirri  shorter,  about  twice  the  length  of  the  following  cirri ;  rest  of  dorsal 
cirri  short,  less  than  half  the  body  width  and  extending  only  slightly 
beyond  the  setae,  subequal.  Short  epaulettes  (a  pair  of  shallow,  semi- 
circular raised  areas)  on  tentacular  segment  only.  Of  68  specimens  of 
the  stem  form,  45  showed  bud  formation  of  the  sexual  stolons,  the 


248  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  los 

head  of  the  bud  forming  between  setigers  13  and  14;  6  were  broken 
after  setiger  13,  5  consisted  of  13  setigers  plus  a  definite  posterior 
growing  zone,  and  12  consisted  of  36  setigers  or  less  and  showed  no 
sign  of  bud  formation.  Of  the  45  specimens  showing  bud  formation, 
18  were  young  buds  (sex?)  showing  four  eyes  and  rudiments  of  head 
appendages;  17  were  developing  male  stolons  as  indicated  especially 
by  the  developing  forked  palps  (at  first,  one  part  of  fork  about  one- 
half  the  length  of  the  other),  large  median  antenna,  and  large  dorsal 
tentacular  cirri;  the  male  buds  consisted  of  a  variable  number  of 
setigers  (14-26  plus  a  definite  growing  zone,  or  32-42  and  tapered 
gradually  to  an  attenuated  posterior  end);  10  were  developing  female 
buds  as  indicated  by  the  developing  three  subequal  antennae  and  the 
absence  of  forked  palps;  the  female  buds  consisted  of  40-65  setigers. 
Color:  In  alcohol:  Two  dorsolateral,  longitudinal  dark  bands  just 
dorsal  to  bases  of  dorsal  cirri  (almost  all  specimens  showed  at  least 
faint  bands).  In  life:  Anterior  part  pale  yellowish  with  dark  dorso- 
lateral bands,  posterior  part  orange.  Tubes  :  Thin,  tough,  transparent, 
on  hydroids  as  Lajoeina  maxima  Levinsen;  either  on  the  surface  or 
surrounded  by  the  sessile  hydrothecae  of  the  hydroid  colony. 

Female  stolon  (Sacconereis) :  Length  5-15  mm.,  width  without 
setae  0.5-1  mm.,  width  including  swimming  setae  3.5-4  mm.,  up  to 
26-51  segm^ents.  Body  widest  in  median  region,  tapering  slightly 
anteriorly  and  more  so  posteriorly,  forming  an  attenuated  tail  region; 
divided  into  three  regions  composed  of  6  anterior  unmodified  setigers, 
13-18  modified  setigers  with  long,  iridescent  swimming  setae,  and 
3-27  posterior  unmodified  setigers.  Prostomium  short,  broad,  one 
pair  eyes  dorsolateral,  one  pair  larger  ones  ventrolateral,  both  with 
lenses;  antennae  long,  stout,  subequal  or  median  one  slightly  longer 
than  lateral  pair.  Upper  two  pairs  tentacular  cirri  similar  to  dorsal 
chri  of  following  segments,  with  a  third  pair  of  short,  ventral,  tentacu- 
lar cirri  (about  length  of  prostomium)  and  a  pair  of  small,  ventral, 
achaetous  lobes.  Dorsal  cirri  about  as  long  as  body  width.  Pair 
of  shallow  semicircular  epaulettes  on  tentacular  segment  only.  Pyri- 
form-shaped  egg  sac  carried  on  ventral  surface  in  region  of  setigers 
10-18,  the  egg  mass  flattened  or  slightly  concave  on  side  toward  body, 
the  narrower  end  of  lobe  anterior,  the  eggs  orange-colored  (in  life). 
Color:  In  alcohol:  Colorless,  transparent,  may  be  dusky  posteriorly 
or  somewhat  dusky  throughout.  Two  female  stolons  with  eggs  in- 
side body  dredged  in  75  fms.  October  11,  1949;  females  with  egg  sacs 
taken  from  vertical  plankton  hauls  in  13  fathoms  March  29,  April 
15,  May  17,  1950;  female  with  egg  sac  containing  developing  embryos 
dredged  in  5  fathoms  January  25,  1950. 

Male  stolons  (Polybostrichus) :  Length  7-9  mm.,  width  without 
setae  0.8-1  mm.,  width  including  swimming  setae  2  mm.,  43-66  seg- 


MARINE    POLYCHAETE    WORMS — PETTIBONE  249 

ments,  with  body  regions  of  6  anterior,  27-30  middle,  9-10  posterior 
setigers,  tapering  gradually  posteriorly.  Prostomium  short,  wide, 
with  a  pair  of  eyes  dorsolateral,  a  larger  pan-  ventrolateral,  all  with 
lenses;  palps  bifurcated  about  basal  third,  the  two  branches  subequal; 
very  long,  stout  median  antenna  and  very  small  lateral  antennae 
anterior  and  medial  to  dorsal  pair  of  eyes.  Three  pairs  tentacular 
cirri  with  anterior  dorsal  pair  similar  to  dorsal  cirri,  anterior  ventral 
pair  shorter  than  prostomial  length,  posterior  dorsal  pair  very  large, 
similar  to  median  antenna,  with  a  pair  of  achaetous  knobs  at  their 
ventral  bases.  Dorsal  cirri  of  anterior  setigers  about  as  long  as  body 
width,  those  of  modified  region  about  as  long  as  the  elongated,  paddle- 
like parapodial  lobes,  those  of  posterior,  taillike  region  short,  digitiform. 
Shallow  semicircular  epaulettes  on  tentacular  segment  only.  Color: 
In  alcohol:  Rusty  brown,  darkest  on  anterior  part  or  colorless  except 
anterior  part.  In  life :  Yellowish  green.  Taken  in  plankton,  near  shore, 
September  10,  1949;  dredged  in  27  fathoms  February  18,  1950. 

Remarks. — As  referred  to  under  the  systematic  discussion  of 
Autolytus,  the  stem  form  of  A.  jallax  has  been  incorrectly  referred  to 
A.'proliferhj KugmQv  (1913,  1928)  and  Wesenberg-Lund  (1947,  1950). 
The  two  may  be  distinguished  by  the  different  positions  of  the  sexual 
buds,  the  greater  length  of  the  dorsal  cirri  in  A.  prolifer,  and  the  color 
markings.  The  sexual  stolons  of  A.  jallax  have  been  confused  with 
A.  prismaticus  (see  discussion  above). 

New  records.— -Arctic  Alaska:  Eluitkak  Pass,  Elson  Lagoon  near 
Point  Barrow,  5-6.6  fms.,  mud,  in  tubes  associated  with  hydroid 
Lafoeina  maxima  Levinsen;  Point  Barrow  base,  washed  ashore,  on 
same  hydroid;  off  Point  Barrow  base,  5-75.5  fms.,  up  to  8  miles 
from  shore,  on  bottoms  of  mud,  stones,  and  various  combinations  of 
mud,  gravel,  stones,  rocks,  large  perforated  rocks,  with  bryozoans, 
hydroids  including  Lafoeina  maxima,  and  shells  (16  stations,  85 
specimens);  sexual  stages  from  plankton  near  shore  to  75.5  fms.  (7 
stations,  22  specimens).  Cape  Smyth,  Point  Barrow  Expedition, 
winter,  1883  (1  male,  2  females  with  egg  sacs).  East  Coast  North 
America:  Off  Labrador,  25-60  fms.,  Blue  Dolphin  Expeditions,  1949, 
1951. 

Distribution. — Widely  distributed  in  the  Arctic:  Alaskan  and 
Canadian  Arctic,  Greenland,  Spitsbergen,  Franz  Josef  Land,  Novaya 
Zemlya.  Also  Faroes;  Labrador.  In  low  water  to  75.5  fathoms; 
sexual  stolons  in  plankton. 

Autolytus  prismaticus  (Fabricius,  1780) 

Figure  29,  a,  b 

Nereis  prismatica  ?  MuUer,  1776,  p.  218.— Fabricius,  1780,  p.  302;  1799,  p.  177, 

pi.  4,  figs.  17-20. 
?  Polybostrichus  longosetosus  Oersted,  1843,  p.  183,  pi.  5,  figs.  62,  67,  71. 


250  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  103 

?  Autolytus  longisetosus  Malmgren,  1867,  p.  34,  pi.  7,  fig.  38. — Verrill,  1881,  p. 

292,  pi.  12,  fig.  10.— Moore,  1902,  p.  274;  1909b,  p.  134. 
Autolytus  incertus  Malmgren,  1867,  p.  35,  pi.  6,  fig.  40. 
Proceraea  gracilis   Verrill,  1874a,  pp.  43,  132,  pi.  5,  fig.  1;  1874b,  pp.  361,  362, 

370,  379,  pi.  3,  fig.  2.— Webster  and  Benedict,  1887,  p.  723. 
Autolytus  prismaticus  Marenzeller,   1892,  p.  420. — Chamberlin,   1920,  p.   12. — 

Augener,    1928,   p.    725.— Annenkova,    1934,   p.   322;    1938,   p.    156.— Not 

Berkeley  and  Berkeley,  1938,  p.  48;  1948,  p.  68.— Gorbunov,  1946,  p.  38.— 

Wesenberg-Lund,  1947,  p.  24,  figs.  10-12  (part). —  ?  Hartman  and  Reish, 

1950,  p.  13. 
Autolytus  trilineatv^  Berkeley  and  Berkeley,  1945,  p.  318,  fig.  1,  a-b;  1948,  p. 

69,  fig.  100. 

Description.- — ^Stem  form:  Length  5-24  mm.,  mdth  without  setae 
0.7-1  mm.,  up  to  52-92  segments.  Proportion  of  anterior  appendages 
and  dorsal  cirri  similar  to  A.  fallax.  Distinct  epaulettes  on  tentacular 
segment  and  usually  extending  also  on  at  least  part  of  first  setigerous 
segment.  Of  7  specimens,  2  were  without  signs  of  bud  formation, 
although  the  body  was  constricted  more  than  normally  between 
setigers  13  and  14;  4  showed  sexual  bud  formation  with  the  head 
developing  between  setigers  13  and  14;  and  1  had  13  setigers  mth  a 
newly  regenerating  posterior  end.  Color:  In  alcohol:  Creamy  white 
with  three  conspicuous,  longitudinal,  black  or  dark  bands,  the  median 
one  broader,  less  dense,  continuous  throughout  body,  the  dorsolateral 
ones  narrower,  darker,  at  level  of  bases  of  dorsal  cirri,  often  confined 
to  anterior  fourth  or  half  of  body. 

Female  stolon  (Sacconereis) :  A  specimen  of  58  setigers,  9  mm.  long, 
1  mm.  wide  without  setae,  3  mm.  wide  with  swimming  setae,  with 
body  regions  of  6  anterior,  18  middle,  and  34  posterior  setigers. 
Epaulettes  somewhat  triangular,  extending  on  tentacular  and  first 
setigerous  segments.  Otherwise  SiS  in  A.  fallax.  Color:  In  alcohol: 
Shows  the  same  characteristic  pigmentation  as  the  stem  form,  three 
longitudinal,  black  pigmented  bands.  Two  female  stolons  with 
swimming  setae  and  eggs  massed  inside  body  dredged  in  5  and  6 
fathoms  August  6,  30,  1948.  Male  stolons  {Polyhosirichus) :  no 
specimens  taken. 

Remarks. — Polyhosirichus  longosetosus  Oersted,  the  male  stolon 
(this  might  well  be  the  male  stolon  of  A.  fallax,  see  systematic  dis- 
cussion above),  Autolytus  incertus  Malmgren,  the  female  stolon,  and 
Proceraea  gracilis  Verrill,  the  stem  form,  have  been  referred  previously 
by  Marenzeller  (1892)  to  A.  prismaticus.  A.  trilineatus  Berkeley  and 
Berkeley  is  herein  referred  to  A.  prismaticus]  the  description,  including 
the  characteristic  three  longitudinal  bands,  is  in  agreement.  The 
record  of  A.  prismaticus  by  the  Berkeleys  (1938,  1948)  is  doubted 
since  the  stem  form  agrees  with  A.  cornutus  or  A.  prolifer  but  not  A. 
prismaticus;  the  sexual  forms  agree  with  A.  cornutus  or  A.  prismaticus 
in  the  number  of  six  anterior  unmodified  setigers;  they  agree  with  A. 


MARINE    POLYCHAETE    WORMS — ^PETTIBONE 


251 


^frkn  ,p 


tCi"' 


Figure  29. — Syllidae:  a,  Autolytus  prismaticus,  stem  form,  dorsal  view  anterior  end;  h 
same,  parapodium;  c,  Autolytus  fallax,  female  (Sacconereis)  form,  dorsal  view  anterior 
end;  d,  same,  ventral  view  anterior  end;  e,  Autolytus  fallax,  male  (Polybostrichus)  form, 
dorsal  view  anterior  end;  /,  same,  ventral  view  anterior  end.  (For  explanation  of  symbols, 
see  p.  210.) 


261112—54- 


252  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  103 

comutus  as  to  size,  about  one-third  the  size  of  the  sexual  stage  of  A. 
prismaticus,  and  in  the  absence  of  semicircular  epaulettes  (specimens 
examined  through  the  kindness  of  the  Berkeleys).  The  record  of  A. 
prismaticus  from  Oregon  by  Hartman  and  Reish  (1950)  is  doubted 
since  it  is  based  on  male  stolons  only. 

New  records. — Arctic  Alaska:  Eluitkak  Pass,  Elson  Lagoon  near 
Point  Barrow,  6.6  fms.,  stony-mud;  off  Point  Barrow  base,  5-70  fms., 
up  to  7  miles  from  shore,  on  bottoms  of  stones,  and  various  combina- 
tions of  mud,  gravel,  stones  (7  stations,  11  specimens).  East  Coast 
North  America:  Lagoon  Pond,  Martha's  Vineyard,  Massachusetts, 
pile  scrapings,  Pettibone,  1951. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian,  Alaskan, 
and  Canadian  Arctic,  Greenland,  Spitsbergen,  Barents  Sea,  Novaya 
Zemlya.  Also  Iceland;  Labrador  to  Massachusetts;  Bering  Sea  to 
British  Columbia  ( ?  Oregon) ;  north  Japan  Sea.  In  low  water  to  267 
fathoms;  sexual  forms  at  surface. 

Genus  Syllis  Savigny,  1818 

Both  species  represented  have  the  body  elongate,  slender,  with 
numerous  segments,  flattened  ventrally,  arched  dorsally,  tapering 
slightly  anteriorly  and  gradually  posteriorly.  Prostomium  oval,  wider 
than  long;  two  palps  wider  basally,  well  separated  except  where  their 
inner  basal  sides  are  fused  (fig.  28,  c).  Dorsal  cirri  distinctly  monili- 
form  throughout  body,  alternately  slightly  longer  and  shorter.  Ven- 
tral cirri  digitiform,  slightly  shorter  or  longer  than  the  parapodial 
lobes  (fig.  28,  d).  Anterior  end  of  extended  proboscis  with  a  smooth, 
chitinous  rim,  a  single  large,  grayish  dorsal  tooth,  with  a  ring  of  short, 
thick  papillae,  with  a  ring  of  10  shorter  papiUae  more  basally.  Male 
and  female  stolons  or  epitokous  forms  (Chaetosyllis  Malmgren,  1867) 
developed  from  the  posterior  segments  of  the  body;  when  fully 
developed  consisting  of  small  bilobed  (or  tetralobed)  bulbous  "head" 
with  two  pairs  of  eyes,  a  dorsal  pair  and  a  larger  ventral  pair,  with 
a  pair  of  short  moniliform  antennae ;  the  first  setigerous  segment  with 
neurosetae  only,  followed  by  a  variable  number  of  segments  with 
additional  long  capillary  swimming  setae  and  usually  a  few  unmodified 
posterior  segments;  stolons  greatly  distended  with  developing  sex 
products;  neurosetae,  dorsal,  ventral  and  anal  cirri  similar  to  those 
of  stem  form. 

Key  to  the  species  of  Syllis  from  Point  Barrow 

1.  Compound  neurosetae  with  short  and  long  distal  blades  (fig.  28,  /).  Acicula 
of  parapodia  not  particularly  enlarged  or  protruding.  Body  segments  very 
short S.  cornuta 

Compound  neurosetae  with  terminal  blades  differing  very  slightly  in  length 
(fig.  28,  e) .  From  1  to  4  very  large  acicula  usually  protrude  somewhat  from 
parapodial  tip  (fig.  28,  d).    Body  segments  longer S.  fasciata 


MARESTE    POLYCHAETE    WORMS — PETTIBONE  253 

Syllis  (Ehlersia)  cornuta  Rathke,  1843 

Figure  28,  f 

Syllis  (Ehlersia)  cornuta  Rathke,  1843,  p.  164,  pi.  7,  fig.  12. — Malmgren,  1867, 

p.  43,  pi.  7,  fig.  45.— Southern,  1914,  p.  37.— Fauvel,  1923,  p.  267,  fig.  100, 

g-i.— Augener,  1928,  p.  718.— Monro,  1933,  p.  34;  1937,  p.  273;  1939b,  p. 

387.— Wesenberg-Lund,  1947,  p.  6;  1950a,  p.  15;  1950b,  p.  46;  1951,  p.  36.— 

Zatsepin,  1948,  p.  115,  pi.  31,  fig.  4. 
Chaetosyllis  oerstedi  Malmgren,  1867,  p.  45,  pi.  8,  fig.  51  (epitokous  form). 
Syllis  oerstedi  Th6el,  1879,  p.  40,  pi.  2,  fig.  33;  pi.  3,  figs.  34,  35.— Annenkova, 

1938,  p.  150.— Gorbunov,  1946,  p.  38. 
Syllis  quaternaria  Moore,  1906a,  p.  352,  fig.  (epitokous  form). 
Syllis  alternata  Moore,   1908,  p.  321,  fig.  (p.  324);  1909a,  p.  321. — Annenkova, 

1938,  p.  148.— Rioia,  1941,  p.  691,  pi.  3,  figs.  1-9.— Berkeley  and  Berkeley, 

1948,  p.  77,  fig.  115. 
Syllis  (Ehlersia)  heterochaeta  Moore,  1909a,  p.  322,  pi.  15,  figs.  1-4. — Annenkova, 

1938,  p.  148.— Rioja,  1941,  p.  694,  pi.  4,  figs.  7-10;  Berkeley  and  Berkeley, 

1948,  p.  76,  fig.  113. 
Ehlersia  cornuta  Hartman,  1945,  p.  15. 
Typosyllis  alternata  Hartman,  1948,  p.  21. 

Description. — Stem  form:  Length  9-45  mm.,  width  without  setae 
1-1.2  mm.  Segments  very  short.  Prostomium  with  two  pairs  of  eyes, 
anterior  pair  larger,  crescentric,  with  or  without  pair  of  small  ocular 
spots  anterior  to  lateral  antennae.  Moniliform  dorsal  cirri  with  about 
14-22  articles  (11-40).  Neurosetae  all  compound,  with  longer  and 
shorter  blades,  the  longer  ones  2-4  times  longer  than  the  shorter  ones, 
with  jfine  marginal  fringe  extending  to  near  the  tips  (may  give  appear- 
ance of  having  a  bidentate  tip  with  a  subterminal  tooth) ;  tips  of  short 
blades  slightly  hooked;  tips  of  long,  thin  blades  slightly  hooked  or 
with  a  slight  knob  (worn  down?).  In  posterior  segments,  terminal 
blades  of  neurosetae  not  as  long  as  in  anterior  segments.  One  speci- 
men with  developing  stolon  of  41  segments  beginning  on  segment 
75  of  stem  form.  Color:  In  alcohol:  (1)  anterior  segments  dusky, 
then  two  narrow,  reddish  brown,  transverse  bands  per  segment  on 
anterior  and  middle  part  of  each  segment;  (2)  rusty  brown,  with  two 
darker  bands  per  segment  on  part  of  body;  (3)  colorless. 

Male  and  female  stolons  (Chaetosyllis) :  Length  8-19  mm.,  width 
without  setae  1-1.2  mm.,  width  including  setae  4-5  mm.,  segments 
30-70  (1  unmodified  anterior  segment,  24-62  modified  segments  with 
swimming  setae,  0-7  unmodified  posterior  segments);  dorsal  cirri 
alternately  longer  and  shorter,  with  11-18  articles  (14-25 — Moore, 
1906);  capillary  setae  much  longer  than  the  dorsal  cirri;  neurosetae  as 
in  posterior  segments  of  stem  form  (distal  blades  not  as  long  as  in 
anterior  region  of  stem  form) ;  some  segments  showed  the  characteristic 
banding  of  two  narrow  bands  per  segment. 


254  PROCEEDINGS   OF   THE    NATIONAL   MUSEUM  vol.  los 

Remarks. — The  types  of  Syllis  alternata  Moore  from  Alaska  and 
S.  heterochaeta  Moore  from  California  were  examined  and  are  herein 
referred  to  S.  cornuta  (suggested  previously  by  Monro  (1933)  for  S. 
heterochaeta).  The  description  of  S.  quaternaria  Moore  (1906),  the 
epitokous  form  taken  at  the  surface,  Point  Barrow,  Alaska,  agrees 
with  that  of  the  stolons  of  S.  cornuta  collected  by  Dr.  MacGinitie  in 
a  plankton  haul. 

New  records. — Arctic  Alaska:  Stem  form:  Off  Point  Barrow  base, 
18.3-123.5  fms.,  up  to  15  miles  from  shore,  on  bottoms  of  stones,  mass 
of  worm  tubes,  and  various  combinations  of  mud,  pebbles,  stones, 
gravel,  rocks,  large  perforated  rocks,  and  worm  tubes  (17  stations,  97 
specimens).  Sexual  stolons:  Off  Point  Barrow  base,  1.6  miles  from 
shore,  vertical  plankton  haul  of  13  fathoms  tlirough  hole  in  ice,  March 
29,  April  15,  1950  (12  specimens).  West  Coast  North  America: 
Strait  of  Juan  de  Fuca  and  Puget  Sound,  Washington,  30-107  fms., 
Pettibone.  East  Coast  North  America:  Off  Martha's  Vineyard, 
86,  134,  and  146  fms.;  off  Salem,  Massachusetts,  35  fms.,  U.  S.  Fish 
Commission, 

Distribution. — Cosmopolitan.  Widely  distributed  in  the  Arctic: 
Siberian  and  Alaskan  Arctic,  Greenland,  Spitsbergen,  Novaya  Zemlya. 
Also  Iceland,  Norway  to  Madeira,  Mediterranean;  Red  Sea,  South 
Arabian  coast,  Persian  Gulf;  Indian  Ocean  (Zanzibar);  off  South 
Africa;  Maine  to  North  Carolina;  north  Japan  Sea,  Alaska  to  Panamd; 
South  Pacific  (Marquesas,  Tahiti).  In  low  water  to  1,400  fathoms; 
sexual  stolons  in  plankton. 

Syllis  {Typosyllis)  fasciata  Malmgren,  1867 

FiauRE  28,  o-e 

Syllis  (Typosyllis)  fasciata  Malmgren,  1867,  p.  43,  pi.  7,  fig.  47;  pi.  8,  fig.  52. — 
Augener,  1928,  p.  719.— Fauvel,  1934b,  p.  304.— Annenkova,  1934,  p.  322; 
1938,  p.  150.— Gorbunov,  1946,  p.  38.— Wesenberg-Lund,  1947,  p.  10,  fig. 
2,a;  1950a,  p.  16;  1950b,  p.  47;  1951,  p.  37.— Berkeley  and  Berkeley,  1948, 
p.  74,  figs.  109,  110.— Zatsepin,  1948,  p.  115,  pi.  31,  fig.  2. 

Syllis  armillaris  Treadwell,  1937,  p.  28  (not  Nereis  armillaris  Mtiller,  1776;  in 
USNM). 

Description. — Stem  form:  Length  8-75  mm.,  width  without  setae 
0.7-3  mm.  Many  smaU  ones  present;  many  broken  ones  and  a 
number  with  regenerating  posterior  ends.  Segments  not  as  short  as 
in  S.  cornuta.  Prostomium  with  two  pairs  of  eyes,  anterior  pair 
larger,  crescentric.  Moniliform  dorsal  cirri  with  about  24  articles 
(20-40;  12-17  in  young  specimens).  Parapodia  each  with  one  to 
four  large,  pointed  acicula  usually  protruding  beyond  distal  tips  of 
setal  lobes.  Neurosetae  all  compound,  with  terminal  blades  rather 
short,  hooked,  with  tips  entire;  some  setae  may  have  blades  broken 


MARINE    POLYCHAETE    WORMS — PETTIBONE  255 

off  (not  to  be  confused  with  simple  setae).  Color:  In  alcohol:  (1) 
pigmented  with  reddish-brown,  one  wide  band  per  segment  (darker 
on  posterior  part  of  band) ;  more  posteriorly  two  bands  per  segment, 
lighter  anterior  and  darker  posterior  ones;  then  one  band  per  segment; 
then  without  color  (when  mature,  banded  color  pattern  on  the  seg- 
ments of  the  developing  sexual  stolon);  (2)  colorless,  particularly  in 
small  specimens;  (3)  uniform  brown  mottling  on  anterior  third  of 
body  (specimens  from  Washington).  In  life:  Reddish  rusty  brown 
bands,  one  wide  band  or  two  narrower  bands  per  segment,  especially 
on  the  anterior  fourth  and  posterior  fourth  (latter  a  developing  stolon, 
with  wide,  orange-red  bands).  One  specimen  with  developing  stolon 
beginning  on  segment  71. 

Sexual  stolons  (Chaetosyllis) :  Two  male  stolons,  34-37  segments 
(one  unmodified  anterior  segment,  31-33  modified  segments  with 
swimming  setae,  2  or  3  small,  unmodified  posterior  segments);  long 
swimming  notosetae  subequal  to  dorsal  cirri;  dorsal  cirri  of  first 
unmodified  segment  broken  off  at  cirrophores;  darkly  pigmented 
reddish  brown  dorsaUy  and  ventrally. 

New  Records. — Arctic  Alaska:  Stem  form:  Off  Point  Barrow  base, 
18.3-123.5  fms.,  up  to  15  miles  from  shore,  on  bottoms  of  mud,  stones, 
worm  tubes,  from  breaking  rocks  and  bryozoans,  from  interstices 
between  pebbles  and  gravel  covering  tunicate,  Molgula  sp.,  from 
various  combinations  of  mud,  pebbles,  stones,  gravel,  rocks,  large 
perforated  rocks,  worm  tubes,  and  shells  (21  stations,  206  specimens). 
Male  stolons:  Off  Point  Barrow  base,  1.6  miles  from  shore,  vertical 
plankton  haul  of  13  fms.  through  hole  in  ice  (March  29,  1950,  2  speci- 
mens). West  Coast  North  America:  Washington  Sound,  17-21 
fms.,  Pettibone.  Canadian  Arctic:  Center  Foxe  Basin,  25-31  fms., 
Bartlett,  1927. 

Distribution. — Widely  distributed  in  the  Ai'ctic:  Siberian,  Alaskan, 
and  Canadian  Ai-ctic,  Baffin  Bay,  Greenland,  Jan  Mayen,  Spitsbergen, 
Franz  Josef  Land,  Novaya  Zemlya,  Kara  Sea.  Also  Iceland,  Faroes; 
Bering  Sea  to  southern  California;  north  Japan  Sea,  China.  In  low 
water  to  378  fathoms;  sexual  stolons  in  plankton. 

Genus  Sphaerosyllis  Claparfede,  1863 

Represented  by  a  single  species  from  Point  Barrow. 

Sphaerosyllis  erinaceus  Clapar^de,  1863 

Figure  28,  m 

Sphaerosyllis  erinaceus  Claparfede,  1863,  p.  45,  pi.  13,  fig.  38  (fide  Fauvel). — 
Southern,  1914,  p.  20. — Fauvel,  1923,  p.  302,  fig.  115,  q-r. — Annenkova, 
1934,  p.  322;  1938,  p.  153.—  ?  Rioja,  1943,  p.  211,  figs.  1-6.— Gorbunov,  1946, 
p.  38.— Zatsepin,  1948,  p.  116,  pi.  31,  fig.  7. 


256  PROCEEDINGS    OF   THE   NATIONAL   MUSEUM  vol.  103 

Sphaerosyllis  latipalpis  Levinsen,  1882,  p.  244. — Augener,  1928,  p.  722,  pi.  11, 
fig.  3;  1939,  p.  140.— Wesenberg-Lund,  1947,  p.  13,  fig.  4;  1951,  p.  38. 

Sphaerosyllis  longicauda  Webster  and  Benedict,  1887,  p.  720,  pi.  3,  figs.  35-39. — 
Eliason,  1920,  p.  11,  fig.  3. 

Oophylax  minuta  Treadwell,  1937,  p.  29,  figs.  6,  7. 

Brania  sp.  Hartman,  1944a,  pi.  24,  figs.  1,  2. 

Description. — Length  2-4.5  mm.,  width  without  setae  0.3-0.5  mm., 
segments  22-37.  Body  linear,  tapering  slightly  anteriorly  and 
posteriorly,  oval  in  cross  section.  Body,  including  parapodia,  covered 
with  small  papillae  and  incrusted  with  fine,  granular  material.  Pro- 
stomium  subrectangular,  wider  than  long,  with  six  black  eyes,  four 
larger  ones  (each  with  a  lens,  arranged  in  transverse  line  or  slightly 
concave  arc  near  middle  of  prostomium)  and  two  small  anterior  ones. 
Antennae  subequal,  bulbous  basally,  narrower  distally.  Paired 
antennae  lateral  to  anterior  eye  spots;  median  antenna  more  posterior, 
nearly  in  line  with  the  four  larger  eyes.  Palps  short,  wide,  rounded 
anteriorly;  they  may  project  more  anteriorly,  extending  about  the 
same  length  as  the  prostomium,  with  basal  halves  fused  and  distal 
halves  separated  by  a  narrow  groove,  or  they  may  project  more  ven- 
trally,  extending  only  about  half  the  length  of  the  prostomium,  and 
fused  except  for  a  small  anterior  indentation;  ventrally  the  palps 
are  not  fused  although  they  may  be  closely  approximated,  extending 
back  to  the  level  of  the  four  large  eyes.  Tentacular  segment  not 
distinctly  set  off  from  the  prostomium,  with  a  single  pair  of  tentacular 
cirri  which  are  similar  to  the  antennae.  Dorsal  cirri  short,  slightly 
longer  than  parapodial  lobes  and  shorter  than  the  setae,  inflated 
basally,  tapering  to  narrow  tips;  they  may  be  similar  in  shape  through- 
out body  or  they  may  be  only  slightly  inflated  basally  in  middle  and 
posterior  regions.  Dorsal  cirri  lacking  on  setiger  2.  Ventral  cirri 
digitiform,  slightly  shorter  than  parapodial  lobes.  Anal  cutI  larger 
than  dorsal  cirri;  anal  segment  with  several  larger  papillae.  Para- 
podial tip  usually  with  a  larger  papilla.  Neurosetae  consisting  of  a 
single  long,  simple,  tapering,  curved  upper  seta,  the  rest  compound 
with  distal  blades  long,  subequal,  entire,  hooked,  and  finely  pectinate. 
Colorless  in  alcohol.  Female  (21  fathoms,  September  9,  1948)  with 
large,  rounded  eggs  (larger  in  diameter  than  parapodial  length)  fastened 
to  dorsal  surface  rather  irregularly  between  setigers  8-24,  one  to  four 
per  segment,  attached  between  neuropodia  and  dorsal  cirri  and  just 
medial  to  the  dorsal  cirri;  another  female  (36  fathoms,  October  6, 
1949)  with  large,  oval-shaped  developing  larvae  (developing  setae 
visible)  attached  to  the  dorsal  surface;  swimming  setae  lacking  (with 
swimming  setae  in  male — Fauvel,  1923;  swimming  setae  m  both  male 
and  female  beginning  on  setiger  8 — Webster  and  Benedict,  1887; 
eggs  on  ventral  base  of  feet — Wesenberg-Lund,  1947). 


MARINE    POLYCHAETE    WORMS — PETTIBONE  257 

Remarks. — The  type  of  Oophylax  minuta  Treadwell  from  Foxe 
Channel  was  examined  and  is  herein  referred  to  Sphaerosyllis  erinaceus; 
although  not  mentioned  in  the  original  description,  the  specimen  has  a 
median  antenna  (seen  with  difficulty  on  the  prepared  slide);  neuro- 
setae  include  a  simple  seta  in  addition  to  the  compound  ones,  setiger 
2  lacks  dorsal  cirri.  The  type  specimens  of  S.  longicauda  Webster  and 
Benedict  from  Eastport,  Maine,  were  examined  also  but  they  are 
unsatisfactory,  being  hardened,  shrunken,  and  covered  with  crystals; 
the  description  is  fairly  complete  and  agrees  with  S.  erinaceus  as 
does  the  description  by  Augener  of  S.  latipalpis  Levinsen.  The 
record  by  Kioja  of  S.  erinaceus  from  Mexico  is  questioned  chiefly 
because  of  the  position  of  the  eyes. 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  16.7-75.5 
fms.,  up  to  8  miles  from  shore,  on  bottoms  of  stones  and  various 
combinations  of  gravel,  stones,  rocks,  large  perforated  rocks,  with 
bryozoans  and  hydroids  (8  stations,  37  specimens).  East  Coast 
North  America:  Off  Labrador,  25  fms..  Blue  Dolphin  Expedition, 
1949.  Vineyard  Sound,  Massachusetts,  surface,  U.  S.  Fish  Commis- 
sion, August  3,  1881. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian  ,  Alaskan, 
and  Canadian  Arctic,  West  Greenland,  Spitsbergen.  Also  Iceland, 
Denmark,  Ireland,  English  Channel,  Baltic;  Labrador  to  Mas- 
sachusetts; Bering  Sea;  north  Japan  Sea;  ?  Mexico.  In  low  water 
to  75.5  fathoms;  sexual  forms  at  surface. 

Genus  Exogone  Oersted,  1845 

Both  species  represented  are  very  small,  slender,  linear-elongate, 
tapered  slightly  anteriorly  and  more  so  posteriorly,  flattened  ventrally, 
arched  dorsally.  Body  colorless,  translucent  (in  alcohol).  Pro- 
stomium  oval  to  subrectangular,  arched  somewhat  anteriorly,  much 
wider  than  long;  four  eyes,  with  lenses,  in  trapezoidal  arrangement; 
palps  fused  dorsally  into  a  rounded  or  triangular  mass,  with  or  without 
a  slight  anterior  notch;  palps  separated  ventrally  by  wide  depression. 
Tentacular  segment  short,  more  or  less  distinct,  the  anterior  part  of 
which  may  form  a  fold  covering  the  posterior  part  of  the  prostomium. 
Dorsal  cirri  shorter  than  parapodial  lobes,  ovoid  to  somewhat  conical. 
Ventral  cirri  digitiform,  shorter  than  parapodial  lobes.  Neurosetae  of 
three  kinds:  (1)  upper  one  simple,  slightly  curved;  (2)  composite, 
with  longer,  slender,  pointed  blades;  (3)  composite,  with  short,  hooked 
blades.  Everted  pharynx  crowned  with  circle  of  papillae  and  single 
conical  tooth.  With  swimming  setae  at  maturity.  Large  eggs, 
embryos  or  young  (up  to  five  setigers,  resembling  the  adult)  attached 
to  ventral  surface  of  female. 


258  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol,  103 

Key  to  the  species  of  Exogone  from  Point  Barrow 

I.  Antennae  subequal.     Tentacular  cirri  rudimentary,  reduced  to  small  knobs 

(fig.  28,  0-     Dorsal  cirri  lacking  on  setiger  2.     Upper  compound  setae  with 

rather  short  pointed  blades E.  naidina 

Median  antenna  more  than  4  times  length  of  lateral  antennae  (fig.  28,  k). 
Tentacular  cirri  only  slightly  smaller  than  the  lateral  antennae  or  dorsal 
cirri.  Dorsal  cirri  present  on  all  setigerous  segments.  Upper  compound 
setae  with  long  pointed  blades E.  dispar 

Exogone  naidina  Oersted,  1845 

Figure  28,  I 

Exogone  naidina  Oersted,  1845,  p.  20,  pi.  2. — Augener,  1939,  p.  140. — Zatsepin, 
1948,  p.  116,  pi.  31,  fig.  8. 

Exogone  gemmifera  Pagenstecher,  1862,  p.  267,  pi.  25;  pi.  26,  figs.  1,  2,  6-8. — 
Mcintosh,  1908,  p.  151,  pi.  59,  figs.  5,  6.— Southern,  1914,  p.  17.— Eliason, 
1920,  p.  11.— Fauvel,  1923,  p.  305,  fig.  117,  a-d.— Annenkova,  1934,  p.  322; 
1938,  p.  154.— Rioja,  1943,  p.  223,  figs.  38-46.— Gorbunov,  1946,  p.  38.— 
Thorson,  1946,  p.  35,  fig.  8.— Berkeley  and  Berkeley,  1948,  p.  79,  fig.  118.— 
Hartman  and  Reish,  1950,  p.  13. 

Description. — Length  3.5-5  mm.,  width  without  setae  0.3  mm., 
segments  32-36.  Palps  form  rounded  mass,  1.4-2  times  length  of 
prostomium.  Antennae  shorter  than  palps,  subequal  or  median  one 
slightly  longer,  clavate,  inserted  anterior  to  eyes  in  almost  a  straight 
line.  Tentacular  cirri  very  small,  reduced  to  small  knobs.  Dorsal 
cirri  lacking  on  setiger  2.  Neurosetae  of  each  parapodium  consisting 
of  one  simple,  cm-ved  upper  seta,  one  upper  composite  seta  with 
longer  pointed  blade,  and  four  or  five  composite  setae  with  short, 
hooked  blades.  Female  (21  fathoms,  September  15,  1948)  without 
special  swimming  setae,  with  large  eggs  attached  to  ventral  surface 
(between  segments  1 1-25,  one  to  three  per  segment)  medial  to  ventral 
cirri  and  between  parapodia.     Another  female  (57  fathoms,  October 

II,  1949)  with  eggs  on  ventral  surface  of  segments  13-21  (mostly 
four  per  segment) ;  eggs  large,  nearly  touching.  (Swimming  setae  on 
females  with  internal  ova,  absent  in  females  with  ova  attached  to 
ventral  surface,  two  eggs  per  segment — Thorson,  1946.) 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  16.7-123.5 
fms.,  up  to  16  miles  from  shore,  on  bottoms  of  rocks,  stones,  worm 
tubes,  and  various  combinations  of  mud,  gravel,  stones,  rocks,  large 
perforated  rocks,  with  worm  tubes,  bryozoans,  hydroids;  on  female 
Hyas  coarctatus  alutaceus  (14  stations,  82  specimens). 

Distribution. — Alaskan  and  Siberian  Ai-ctic;  Denmark,  Ireland, 
Great  Britain  to  Madeira,  Mediterranean;  Bering  Sea  to  Mexico; 
north  Japan  Sea.  In  low  water  to  123.5  fathoms;  sexual  stages  at 
surface. 


MARINE    POLYCHAETE    WORMS — PETTIBONE  259 

Exogone  dispar  (Webster,  1879) 

Figure  28,  k 

Paedophylax  dispar  Webster,  1879a,  p.  223,  pi.  4,  fig.  49;  pi.  5,  figs.  50-55;  1879b, 

p.  110;  1886,  p.  138.— Sumner,  1913,  p.  615. 
Paedophylax  longiceps  Verrill,  1879,  p.  170;  1881,  p.  320,  pi.  12,  fig.  2;  1882,  p. 

370.— Andrews,  1891,  p.  281. 
Paedophylax  longicirris  Webster  and  Benedict,  1887,  p.  722,  pi.  3,  figs.  46-50. 
Exogone  lourei  Berkeley  and  Berkeley,  1938,  p.  44,  figs.  6-12;  1948,  p.  79,  fig.  117.— 

Rioja,  1941,  p.  703,  pi.  3,  figs.  14-21;  1943,  p.  224. 
Exogone  dispar  Hartman,  1942a,  p.  11;  1944a,  p.  338,  pi.  24,  fig.  9;  pi.  25,  fig.  5; 

1945,  p.  16,  pi.  2,  figs.  7,  9,  10;  1951,  p.  40. 

Description. — Anterior  ends  of  3  specimens  only  (up  to  8  mm.  long, 
45  setigers — Berkeley  and  Berkeley,  1938).  Palps  form  a  conical 
mass,  about  twice  length  of  prostomium.  Antennae  inserted  between 
and  slightly  in  front  of  eyes;  median  antenna  longer,  usually  reaching 
nearly  to  end  of  palps  (may  be  not  much  longer  than  prostomium), 
lateral  antennae  short,  ovoid.  Tentacular  cirri  slightly  smaller  than 
lateral  antennae  and  dorsal  cirri.  Dorsal  cirri  present  on  all  setigerous 
segments,  including  setiger  2,  Neurosetae  on  each  parapodium  con- 
sisting of  one  simple  curved  upper  seta  (may  have  a  delicate  thread 
coming  off  subterminally  in  some  specimens),  one  to  four  composite 
setae  with  long  terminal  capillary  blades,  and  three  to  nine  composite 
setae  with  short,  hooked  blades. 

Remarks. — The  specimens  were  compared  with  the  cotypes  of 
Paedophylax  dispar  from  New  Jersey.  The  types  of  P.  longicirris 
Webster  and  Benedict  from  Eastport,  Maine,  were  examined  also 
and  the  peculiar  hairlike  process  on  some  of  the  simple  setae  was 
noted.  This  also  occurs  on  some  of  the  simple  setae  of  the  specimens 
from  Point  Barrow.  P.  longiceps  Verrill  has  previously  been  referred 
to  P.  dispar  by  Sumner  (1913).  The  description  of  E.  lourei  Berkeley 
and  Berkeley  from  British  Columbia  agrees  with  that  of  E.  dispar. 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  36-70  fms., 
up  to  7.5  miles  from  shore,  on  bottoms  of  stones,  gravel,  large  per- 
forated rocks  (2  stations,  3  specimens) .  West  Coast  North  America: 
Strait  of  Juan  de  Fuca,  Washington,  shore,  Pettibone,  1940.  East 
Coast  North  America:  Woods  Hole  region,  Massachusetts,  Petti- 
bone, 1950,  1951.  Maine,  Massachusetts,  New  Jersey,  low  water  to 
14  fms.,  U.  S,  Fish  Commission. 

Distribution. — Arctic  Alaska  to  Mexico;  Maine  to  southwestern 
Florida.    In  low  water  to  70  fathoms;  sexual  stages  at  surface. 

Genus  Eusyllis  Malmgren,  1867 

Both  species  represented  are  flattened  ventrally,  arched  dorsally, 
tapering  both  anteriorly  and  posteriorly,  very  fragile,  breaking  up 


260  PROCEEDINGS    OF    THE    NATIONAL   MUSEUM  vol.  103 

easily  and  losing  their  antennae  and  dorsal  cirri  readily.  Prostomium 
subrectangular  to  suboval,  wider  than  long;  four  eyes  fairly  large,  in 
trapezoidal  arrangement,  the  anterior  pair  larger;  palps  broad,  thick, 
oval  to  squarish,  fused  at  their  bases  only  and  well  separated  distally, 
and  may  be  curled  ventrally  or  longitudinally  like  flopping  ears; 
lateral  antennae  about  1.5  times  the  prostomial  length;  median 
antenna  about  twice  the  length  of  the  lateral  antennae  (fig.  28,  g). 
Upper  tentacular  cirri  as  long  as  median  antenna,  ventral  pair  much 
shorter.  Ventral  cirri  thick,  oval,  about  same  size  and  length  as 
parapodial  lobes  or  may  be  slightly  shorter  (first  pair  not  enlarged, 
similar  to  the  following).  Neuropodia  short  and  plump,  with  neuro- 
setae  all  composite  except  on  the  last  setigers  where  there  may  be  a 
simple,  entire,  upper  seta  and  a  simple,  bidentate,  lower  seta;  blades  of 
composite  setae  rather  short,  hooked,  distinctly  bidentate,  finely 
spinous  on  the  cutting  edge;  shaft  distally  spinous  (fig.  28,  i).  At 
maturity,  with  long  capillary  swimming  setae.  Proboscis  with  two 
rows  of  soft  papillae,  a  proximal  and  a  distal  row  of  10  papillae  per 
row;  inside  the  distal  row,  a  chitinous  lining  provided  with  a  large, 
triangular,  dorsal  tooth  and  a  denticled,  circular  margin  (fig.  28,  g,  h). 

Key  to  the  species  of  Eusyllis  from  Point  Barrow 

1.  Cirrophores  of  dorsal  cirri  not  prominent.  Tentacular  segment  with  a  wide 
crescent-shaped  nuchal  lobe  covering  the  posterior  part  of  the  prostomium 

(fig.  28,  g) E.  blotnstrandi 

Cirrophores  of  dorsal  cirri  prominent,  as  long  as  the  neuropodia.    Tentacular 
segment  short,  without  nuchal  lobe E.  magnifica 

Eusyllis  blomstrandi  Malmgren,  1867 

Figure  28,  g-i 

Eusyllis  blomstrandi  Malmgren,  1867,  p.  40,  pi.  6,  fig.  43. — Fauvel,  1923,  p.  293, 
fig.  112,h-m.— Augener,  1928,  p.  721.— Annenkova,  1938,  p.  153,  fig.  6.— 
Wesenberg-Lund,  1947,  p.  11,  fig.  3;  1950a,  p.  16;  1950b,  p.  48;  1951,  p.  37.— 
Berkeley  and  Berkeley,  1948,  p.  84,  fig.  126. 

Eusyllis  monilicornis  Malmgren,  1867,  p.  41,  pi.  6,  fig.  44. — Verrill,  1881,  p.  319. — 
Zatsepin,  1948,  p.  116,  pi.  31,  fig.  6,b. 

?  Eusyllis  phosphorea  Hartman,  1942a,  p.  7;  1944a,  pp.  334,  338,  pi.  25,  fig.  4. 

Syllis  monilicornis  Th6el,  1879,  p.  41,  pi.  3,  fig.  39. 

Eusyllis  tubifex  Mcintosh,  1908,  p.  173,  figs.— Southern,  1914,  p.  32  (not  Syllis 
tubifex  Gosse,  1855). 

Eusyllis  bloemstrandi  Friedrich,  1939,  p.  122. 

Typosyllis  collaris  Hartman,  1948,  p.  23,  fig.  6,a-c. 

Description. — Length  7-32  mm.,  width  without  setae  0.8-1.2  mm., 
segments  50-124.  Antennae,  tentacular  cirri,  anterior  dorsal  cirri, 
and  long  anal  cirri  irregularly  annulated,  especially  distally,  more  or 
less  smooth  basally;  the  more  posterior  dorsal  cirri  only  indistinctly 
annulated  or  smooth.  First  few  dorsal  cirri  may  be  longer  than  body 
width,  especially  the  first  pair  which  may  be  the  longest  appendage; 


MARINE    POLYCHAETE    WORMS — PETTIBONE  261 

rest  of  dorsal  cii-ri  shorter  thaii  body  width,  tapering  gradually. 
Tentacular  segment  with  wide,  short,  crescent-shaped  collar  or  nuchal 
fold  covering  posterior  part  of  prostomium,  may  be  inconspicuous 
and  protruding  or  rather  low,  somewhat  contracted.  Composite 
setae  similar  throughout,  with  distal  blades  short,  difTering  only 
slightly  in  length.  Chitinous  lining  of  proboscis  light  or  dark  gray 
or  light  amber-colored;  dorsal  tooth  dark  bluish  gray;  circular  margin 
finely  denticled.  Epitokous  female  (Elson  Lagoon,  August  6,  1948) 
with  large  eggs  massed  in  most  of  body  segments,  consisting  of  16 
anterior  unmodified  setigers,  25  setigers  with  long  swimming  setae, 
and  5  unmodified  setigers  (swimming  setae  begin  on  setiger  13 — 
Wesenberg-Lund,  1947).  Color:  In  alcohol:  Colorless  or  tannish, 
slightly  greenish  anteriorly.  In  life:  Orange  or  yellowish.  May  form 
definite  hardened  mucous  tubes.     Luminescent. 

Remarks. — The  type  of  Typosyllis  coUaris  Hartman  (1948)  was 
examined  and  is  herein  referred  to  E.  hlomstrandi;  the  dorsal  cirri 
are  not  distinctly  annulated  throughout  the  body  as  they  would  be 
in  a  Typosyllis.  There  is  nothing  in  Verrill's  description  of  E.  phos- 
phorea,  dredged  from  St.  George's  Bank,  to  distinguish  it  from  E. 
hlomstrandi;  according  to  Hartman  (1942,  p.  7)  the  type  is  unsatis- 
factory. 

New  records. — Arctic  Alaska:  Eluitkak  Pass,  Elson  Lagoon  near 
Point  Barrow,  6.6  fms.;  Point  Barrow  base,  washed  ashore;  off  Point 
Barrow  base,  13.3-123.5  fms.,  up  to  15  miles  from  shore,  on  bottoms 
of  stones,  masses  of  worm  tubes,  and  various  combinations  of  mud, 
pebbles,  gravel,  stones,  rocks,  large  perforated  rocks,  with  worm  tubes, 
bryozoans,  hydroids,  shells,  and  on  female  crab  Hyas  coarctatus  alu- 
taceus  (24  stations,  175  specimens).  Bering  Sea:  Eobert  White, 
1879.  West  Coast  North  America :  Washmgton  and  Puget  Sounds, 
20-46  fms.,  Pettibone.  East  Coast  North  America:  Off  Labrador, 
15-30  fms..  Blue  Dolphin  Expeditions,  1949,  1951. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian  and 
Alaskan  Arctic,  West  Greenland,  Spitsbergen,  Franz  Josef  Land, 
Barents  Sea,  Novaya  Zemlya,  Kara  Sea.  Also  Iceland,  Faroes,  Ire- 
land to  Madeira,  Mediterranean;  Labrador  to  Maine;  Bering  Sea  to 
Wasliington;  north  Japan  Sea.  In  low  water  to  444  fathoms;  epitokes 
at  surface. 

Eusyllis  magnifica  (Moore,  1906) 

Pionosyllis  magnifica  Moore,  1906b,  p.  223,  pi.  10,  figs.  9-11;  1908,  p.  325.— 
Annenkova,  1938,  p.  152. 

Description.— Atokous  specimen  52  mm.  long,  2.5  mm.  in  greatest 
width  without  setae,  152  segments  (in  pieces,  complete?).  Epitokous 
specimen  (18  fathoms,  September  16,  1948)  broken  up  and  not  quite 
complete,  with  larger  eyes,  consisting  of  161  segments,  99  mm.  long. 


262  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  i03 

4  mm.  wide  without  setae,  9  mm.  wide  including  long  swimming  setae. 
Two  atokous  specimens  from  Washington  43-48  mm.  long,  1.5-2  mm. 
wide  without  setae.  Median  antenna,  tentacular  cirri  and  some  of 
anterior  dorsal  cirri  somewhat  annulated,  especially  distaUy;  rest  of 
dorsal  cirri  smooth.  Some  of  anterior  dorsal  cirri  longer  than  body 
width;  rest  shorter  than  body  width,  subulate,  thick  basally,  tapering 
distally,  on  prominent  cirrophores  which  may  be  as  long  as  parapodial 
lobes.  Tentacular  segment  very  short,  without  nuchal  collar.  Blades 
of  neurosetae  all  rather  short.  In  anterior  parapodia,  blades  of  upper 
neurosetae  somewhat  longer  than  the  lower  ones;  on  rest  of  parapodia, 
blades  of  neurosetae  only  slightly  different  in  length.  Chitinous  lining 
of  proboscis  and  dorsal  tooth  light  to  darker  amber-colored  or  trans- 
parent; circular  anterior  border  with  coarse,  triangular  denticles  (when 
transparent,  difficult  to  see  denticled  border).  Swimming  setae  begin 
on  setiger  24  (on  setiger  19  in  specimen  from  Portage  Bay,  Alaska). 
Color  :  In  alcohol :  Flesh. 

Remarks. — The  specimens  were  compared  with  the  type  of  Piono- 
syllis  magnifica  Moore  (1906)  from  Port  Townsend,  Washington. 
It  is  herein  referred  to  Eusyllis  since  the  composite  neurosetae  have 
rather  short  blades  which  differ  only  slightly  in  length,  the  proboscis 
has  a  proximal  and  a  distal  row  of  papillae,  and  the  anterior  circular 
border  is  denticled  (difficult  to  see  when  the  chitinous  lining  is 
transparent) . 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  18.3  fms., 
pebbles-mud  (1  station,  2  specimens).  West  Coast  North  America: 
Albatross  surface  station.  Portage  Bay,  Alaska;  Washington  Sound, 
Pettibone. 

Distribviion. — Arctic  Alaska  to  Washington;  north  Japan  Sea.  In 
16  to  302.8  fathoms;  epitokes  at  surface. 

Genus  Pionosyllis  Malmgren,  1867 
Pionosyllis  compacta  Malmgren,  1867 

Figure  28,  j 

Pionosyllis  compacta  Malmgren,  1867,  p.  40,  fig.  48. — Augener,  1928,  p.  722. — 
Annenkova,  1934,  p.  322. 

Description. — Anterior  ends  only  of  two  small  specimens,  up  to 
1.5  mm.  wide  without  setae  (8-10  mm.  long,  1  mm.  wide — Malmgren, 
1867).  Body  flattened  ventraUy,  arched  dorsally,  fragile,  with  seg- 
ments crowded.  Prostomium  subrectangular,  wider  than  long;  four 
eyes  rather  large,  in  trapezoidal  arrangement;  palps  squarish,  fused 
basally,  widely  separated  distally.  Tentacular  segment  without 
nuchal  collar.  Antennae,  tentacular,  and  dorsal  cirri  smooth,  without 
articulations,  filiform  to  rather  thick,  breaking  off  readily;  most  of 


MARINE    POLYCHAETE    WORMS — PETTIBONE  263 

dorsal  cirri  shorter  than  the  body  width.  Ventral  cirri  thick,  pointed, 
almost  as  long  as  parapodial  lobes.  Neurosetae  all  composite,  distal 
blades  distinctly  bidentate,  finely  spinous  along  cutting  edge;  blades 
of  upper  setae  longer,  becoming  gradually  shorter  in  lower  part  of 
bundle  with  upper  ones  about  twice  the  length  of  lower  ones.  Proboscis 
with  a  distal  row  of  10  papillae;  transparent  chitinous  lining  with  a 
translucent  bluish-gray  dorsal  tooth,  with  distal  rim  smooth.  Color: 
In  alcohol:  Irregularly  dotted  with  black  pigment  both  dorsally  and 
ventrally.  (Epitokous  forms  have  capillary  setae  beginning  on  seg- 
ments 12-15 — Malmgren,  1867.) 

New  records. — Arctic  Alaska:  Off  Point  Barrow,  36-78  fms.,  up 
to  15  miles  from  shore,  on  bottoms  of  rocks,  stones,  and  worm  tubes 
(2  stations,  2  specimens). 

Distribution. — Arctic  Alaska;  Spitsbergen;  Bering  Sea.  In  low 
water  to  78  fathoms. 

Family  Nereidae 

Prostomium  distinct,  with  four  eyes,  two  frontal  antennae,  two  bi- 
articulated  palps  (fig.  30,  a).  Four  pairs  tentacular  cirri;  buccal  or 
tentacular  segment  usually  apodous  and  achaetous.  Parapodia 
usually  biramous  (except  first  two  pairs),  with  dorsal  and  ventral 
cylindrical  cirri,  two  or  three  notopodial  ligules  and  one  neuropodial 
ligule  (fig.  30,  6,  c).  Setae  usually  compound  spinigers  and  falcigers 
(fig.  30,  d-g).  Pair  of  anal  cirri.  Proboscis  eversible,  with  pair  of 
terminal,  horny,  falcate  jaws  and  usually  with  horny  denticles  or 
paragnaths  arranged  in  eight  groups  on  two  rings,  a  basal  or  oral 
ring  and  a  distal  or  maxillary  ring  (fig.  30,  h,  i).  At  maturity  usually 
with  a  pelagic  epitokous  or  heteronereis  stage. 

Represented  by  a  single  genus  and  two  species. 

Genus  Nereis  Cuvier,  1817 

Subgenus  Nereis  Linne,  1758 

Both  species  have  the  body  smooth,  elongate,  cylindrical,  at- 
tenuated posteriorly,  with  numerous  segments.  Prostomium  of  the 
typical  nereid  form.  Tentacular  segment  achaetous,  over  twice  the 
length  of  the  following  segments;  tentacular  cirri  short,  fairly  uniform 
in  length,  the  upper  one  only  slightly  longer  than  the  others  (may 
reach  setiger  2).  Parapodia  biramous,  with  two  subequal  noto- 
podial ligules  (remaining  so  throughout  the  body,  the  upper  one  not 
getting  larger  or  longer  than  the  lower  one),  a  single  neuropodial 
ligule  ventral  to  the  conical  setigerous  neuropodial  lobe  (fig.  30,  h,  c). 
Notosetae  homogomph  spinigers  (fig.  30,  g);  in  more  posterior  seg- 
ments with  some  homogomph  falcigers  with  blunt  asymmetrical 
reduced    appendages    (fig.    30,    e).     Neurosetae    homogomph    and 


264  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  103 

heterogomph  spinigers  (fig.  30,  /)  and  heterogomph  falcigers  with 
hooked  end-pieces  (fig.  30,  d).  Proboscis  with  paragnaths  all 
chitinous,  conical,  separated  in  groups,  occurring  on  both  rings 
(fig.  30,  h,  i).  Distal  ring  of  paragnaths:  Area  II  (dorsolateral) — 
oblique  group  of  two  or  three  rows;  area  III  (medio ventral) — small 
transverse  group;  area  IV  (ventrolateral)— arched  mass.  Basal  ring 
of  paragnaths:  Area  V  (mediodorsal) — 0.  At  maturity,  with  epito- 
kous  stage,  eyes  enlarged;  14-19  anterior  prenatatory  setigers; 
modified  natatory  setigers  with  parapodial  lobes  compressed,  flat- 
tened anteroposteriorly,  developing  foliaceous  lamellar  plates,  with 
numerous  paddlelike  swimming  setae;  in  males,  first  seven  dorsal 
cirri  enlarged  and  club-shaped,  first  five  or  six  ventral  cirri  modified, 
and  dorsal  cirri  of  natatory  setigers  crenulate  on  lower  margin;  in 
females,  dorsal  and  ventral  cutI  not  modified  or  crenulate. 

Key  to  the  species  of  Nereis  (Nereis)  from  Point  Barrow 

1.  Parapodial  ligules  short,  thick,  evenly  rounded  (fig.  30,  b).  Body  uniformly 
purplish  or  reddish  brown,  not  banded.  Paragnaths  of  distal  ring:  Area 
I  (mediodorsal) — 2  in  tandem  (rarely  1  or  3);  paragnaths  of  basal  ring: 
Area  VI  (dorsolateral) — 4  in  square  or  cross  (rarely  3  or  5);  areas  VII- 
VIII  (ventral) — 1-2  irregular  rows  of  larger  paragnaths,  vnth.  several  rows 
of  small  ones,  diminishing  in  size  posteriorly  (2-3  irregular  rows  of  subequal 
paragnaths  in  subspecies  occidentalis).     Male  epitokes  with  16  and  females 

with  17-19  prenatatory  setigers N.   (N.)  pelagica 

Parapodial  ligules  triangular  to  conical,  gradually  tapering  to  a  broad  tip 
(fig.  30,  c).  Body  transversely  banded  reddish  brown  or  violet.  Parag- 
naths of  distal  ring:  Area  I — 0  or  1;  paragnaths  of  basal  ring:  Area  VI — 
6-10  or  more  in  oval  mass;  VII-VIII — continuous  row  of  larger  parag- 
naths followed  by  a  wide  band  of  small  subequal  ones  (fig.  30,  hy  i).  Mais 
epitokes  with  14  and  females  with  16  prenatatory  setigers N.  (N.)  zonata 

Nereis  {Nereis)  pelagica  Linne  1758 

Figure  30,  a,  b 

Nereis  pelagica  Linn6,  1758,  p.  654. — Malmgren,  1867,  p.  47,  pi.  5,  fig.  35. — 
Webster  and  Benedict,  1884,  p.  718;  1887,  p.  724.— Moore,  1903,  p.  431.— 
Mcintosh,  1908,  p.  268,  figs.— Sumnsr,  1913,  p.  619.— Southern,  1914,  p. 
80.— Eliason,  1920,  p.  29.— Chamberlin,  1920,  p.  14.— Fauvel,  1923,  p.  336, 
fig.  130,  a-f.— Augener,  1928,  p.  712.— Monro,  1930,  p.  106.— Annenkova, 
1934,  p.  322;  1937,  p.  162;  1938,  p.  158.— Treadwell,  1937,  p.  29.— Okuda, 
1939,  p.  230.— Friedrich,  1939,  p.  123.— Hartman,  1940,  p.  225,  pi.  35,  fig. 
52;  1944a,  p.  335;  1948,  p.  26.— Berkeley  and  Berkeley,  1943,  p.  130;  1948, 
p.  66,  fig.  96,  a-b.— Thorson,  1946,  p.  64,  fig.  29.— Zatsepin,  1948,  p.  119, 
pi.  30,  fig.  2.— Wesenberg-Lund,  1950a,  p.  19,  pi.  6,  figs.  26a,  27a;  1950b, 
p.  53;  1951,  p.  39. 

Heteronereis  grandifoUa  Rathke,  1843,  p.  155,  pi.  7,  figs.  13,  14. — Malmgren, 
1865,  p.  108,  pi.  11,  fig.  15  (part — female  heteronereid,  not  male). 

Nereis  neonigripes  Hartman,  1936b,  p.  471,  fig.  48. 

Nereis  arctica  Treadwell,  1937,  p.  30. 

Nereis  pelagica  var.  occidentalis  Hartman,  1945,  p.  20,  pi.  4,  figs.  1-6;  1951,  p.  46. 

Nereis  pelagica  neonigripes  Hartman  and  Reish,  1950,  p.  17. 


MARINE    POLYCHAETE    WORMS — PETTIBONE  265 

Description. — Length  60-155  mm.,  width  5-14  mm.  See  key. 
Color:  In  alcohol:  Iridescent,  uniformly  purplish  or  reddish  brown 
(narrow  white  intersegmental  lines) ;  ligules  of  parapodial  lobes  may 
be  darkly  pigmented  (var.  neonigripes) ;  tips  of  dorsal  cirri  may  be 
black. 

New  records. — ^Arctic  Alaska:  Off  Point  Barrow  base,  up  to  12.1 
miles  from  shore,  18-87  fms.,  on  various  combinations  of  stones, 
gravel,  rocks,  large  perforated  rocks  (8  stations,  10  specimens). 
Spitsbergen:  Spitsbergen  Sea,  U.  S.  S.  Alliance,  August,  1881,  sm-face 
(heteronereid) .  Bering  Sea  :  Albatross  Sta.  at  Nikolski,  Bering  Island, 
1892,  and  Sta.  3496,  56°32'  N.,  169°45'  W.,  41  fms.,  1893;  St.  George 
Island,  Pribilofs,  1913,  1914,  low  tide  and  30  fms.,  and  St.  Paul  Island, 
Pribilofs,  lagoon  reef,  1915,  Hanna;  St.  Paul  Island,  Wm.  Palmer, 
1890.  Southwestern  Alaska:  Albatross  Sta.,  Unalaska,  Aleutians, 
1888;  Akutan  Pass,  Chica  Islands,  1872;  Easka  Harbor,  sandy- 
mud,  9-12  fms.,  1873;  Anchorage,  Big  Koniuji  Islands,  Shu- 
magin,  6-20  fms.,  sand  and  rocks;  Port  Moller,  Alaska  Peninsula, 
beach  to  17  fms.,  sand,  1874;  KiUisnoo,  1897;  Chiniak  Bay,  Kodiak 
Island,  1880;  all  collected  by  Dall.  Gulf  of  Alaska:  Albatross  Sta., 
at  Observation  Island,  Cordova,  Alaska,  1914.  Southeastern 
Alaska:  Sitka,  1932.  West  Coast  North  America:  Albatross 
Sta.  89a,  Denman,  British  Columbia,  1914,  and  Sta.  3466,  off  Wash- 
ington, 48°18'30"  N.,  123°22'  W.,  56  fms.,  1891.  Washington  and 
Puget  Sounds,  low  tide  to  166  fms,  (very  common),  Pettibone.  East 
Coast  North  America:  Off  Labrador,  7-60  fms..  Blue  Dolphin 
Expeditions,  1949,  1951;  Woods  Hole  region,  intertidal  and  dredged, 
Pettibone,  1950,  1951;  Gulf  of  St.  Lawrence,  New  Brunswick,  Nova 
Scotia,  Bay  of  Fundy,  Maine,  Massachusetts,  Ehode  Island,  Con- 
necticut, Long  Island  Sound,  shore  to  250  fathoms  and  surface,  U.  S. 
Fish  Commission. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian,  Alaskan, 
and  Canadian  Arctic,  Davis  Strait,  Greenlaad,  Spitsbergen,  Barents 
Sea,  Novaya  Zemlya,  Kara  Sea.  Also  Iceland,  Faroes,  Norway  to 
Mediterranean;  Hudson  Bay  to  Long  Island  Sound;  Bering  Sea  to 
Panamd;  north  Japan  Sea,  Japan;  South  Atlantic  (Tristan  da  Cunha, 
Kerguelen,  Magellan  Straits).  In  low  water  to  609  fathoms;  epitokes 
at  surface.  Variety  occidentalis  Hartman:  North  Carolina,  Florida, 
Louisiana,  Texas,  Puerto  Rico;  intertidal. 

Nereis  (Nereis)  zonata  Malmgren,  1867 

Figure  30,  c,  h,  i 

Heteronereis  grandifolia  Malmgren,  1865,  p.  108,  pi.  11,  fig.  16  (part — male 
heteronereid,  not  female;  not  H.  grandifolia  Rathke,  1843). 

Nereis  zonata  Malmgren,  1867,  p.  46,  pi.  5,  fig.  34.— Th6el,  1879,  p.  42.— Wir^n, 
1883,  p.  402.— Ditlevsen,  1911,  p.  419,  pi.  28,  fig.  6;  pi.  30,  figs.  18,  22.— 


266  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  103 

Fauvel,  1914,  p.  177,  pi.  14,  figs.  1-17;  1923,  p.  338,  fig.  130,  g-n  (part).— 
Augener,  1928,  p.  713.— Annenkova,  1931,  p.  203;  1932,  p.  134;  1934,  p. 
322;  1937,  p.  162;  1938,  p.  158.— Friedrich,  1939,  p.  123.— Berkeley  and 
Berkeley,  1943,  p.  130.— Gorbunov,  1946,  p.  38.— Zatsepin,  1948,  p.  119, 
pi.  30,  fig.  3. — Hartman,  1948,  p.  25. — Hartman  and  Reish,  1950,  p.  17. — 
Wesenberg-Lund,  1950a,  p.  20,  pi.  6,  figs.  26b,  27b;  1950b,  p.  55;  1951,  p.  41. 

Description. — Length  30-125  mm.,  width  7  mm.  See  key.  Color: 
In  alcohol:  Reddish  or  rusty  brown  or  violet  wide  transvei-se  bands; 
anterior  fourth  of  segment  Avithout  color. 

Remarks. — Nereis  procera  Ehlers,  1868,  originally  described  from 
the  Gulf  of  Georgia,  British  Columbia,  has  been  referred  to  N.  zonata 
by  Fauvel,  1914.  In  A^.  procera  the  hodj  is  long,  slender,  extremely 
attenuated  posteriorly;  the  notopodial  ligules  of  the  anterior  setigers 
are  triangular,  subequal;  those  of  the  median  and  posterior  regions 
have  the  upper  notopodial  ligules  larger  than  the  lower  one  (may  be 
nearly  twice  the  size) ;  the  paragnaths  of  areas  VII  and  VIII  are  in  a 
wide,  continuous  area  with  no  especially  enlarged  anterior  row;  male 
epitokes  with  51  and  females  with  59  prenatatory  setigers.  It  may  be 
that  the  variety  procera  of  Fauvel  refers  to  a  different  species  from  that 
of  N.  procera  Ehlers. 

New  records. — Arctic  Alaska:  Eluitkak  Pass,  Elson  Lagooa  near 
Point  Barrow,  13  fms.,  gravel;  off  Point  Barrow  base,  21-78  fms.,  on 
bottoms  of  gravel,  stones,  small  rocks,  with  shells  and  worm  tubes  (8 
stations,  12  specimens).  Canadian  Arctic:  Ellesmere  Island,  U.S.S. 
Alert,  1950,  J.  Peter  Johnson  (epitoke).  East  Greenland:  74°04'  N., 
17°58'  W.,  120  fms.,  Norcross-Bartlett  Expedition,  1931.  East 
Coast  North  America:  Off  Labrador,  9-100  fms,,  mud.  Blue  Dolphin 
Expeditions,  1950,  1951. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian,  Alaskan, 
and  Canadian  Arctic,  Baffin  Bay,  Davis  Strait,  Greenland,  Norway, 
Spitsbergen,  Franz  Josef  Land,  Barents  Sea,  Novaya  Zemlya,  Kara 
Sea.  Also  Iceland,  Faroes;  Hudson  Bay  to  Labrador;  Bering  Sea  to 
Oregon;  north  Japan  Sea.  In  low  water  to  439  fathoms;  epitokes  at 
surface.  Variety  ?  procera  Fauvel  (1914,  1923;  not  Ehlers,  1868): 
English  Channel  to  Mediterranean.  Variety  persica  Fauvel:  Red 
Sea,  South  Arabian  Coast,  Persian  Gulf,  Indian  Ocean. 

Family  Nephtyidae 

Body  elongated,  subtetragonal  in  cross  section.  Prostomium  small, 
somewhat  flattened,  angulate,  with  four  small  antennae  (fig.  30,  j). 
First  segment  rudimentary,  with  setae  and  usually  with  short  dorsal 
and  ventral  tentacular  chri.  Parapodia  bu-amous,  with  rami  well 
separated,  with  smaU  dorsal  chri  on  lower  sides  of  notopodia;  with 
branchiae  (interramal  chri)  between  the  two  rami,  cirriform,  sickle- 
shaped  or  foliaceous;  with  simple  setae  arranged  in  fan-shaped  bundles; 


MARINE    POLYCHAETE    WORMS — PETTIBONE  267 

ventral  cirri  short,  conical,  or  foliaceous;  with  anterior  and  posterior 
lamellae  more  or  less  developed  (fig.  30,  k-^).  A  single  anal  cirrus. 
Proboscis  eversible,  strong,  muscular,  a  pah-  of  horny  jaws  wdthin  the 
pharynx,  with  rows  of  soft  papillae  and  a  crown  of  terminal,  bifid 
papillae.  Move  rapidly  with  undulating  motion;  usually  in  sand 
or  mud. 

Represented  by  a  single  genus  and  four  species. 

(ienus  Nephtys  Cuvier,  1817 

The  four  species  represented  have  numerous  segments  (65-150), 
tapered  posteriorly  and  slightly  anteriorly.  Prostomium  pentagonal 
to  quadrate,  with  antennae  conical,  subequal,  or  more  ventral  pair 
slightly  larger,  without  eyes,  with  pair  of  nuchal  papillae  at  postectal 
margins  (may  be  inverted  or  everted;  fig.  30,  j).  Tentacular  or  first 
segment  with  both  dorsal  and  ventral  tentacular  cu-ri.  Parapodial 
rami  with  anterior  lamellae  rudimentary  or  lacking,  with  posterior 
lamellae  developed  m  varying  degrees,  with  branchiae  recurved  (con- 
vex side  toward  lateral  side  of  body,  fig.  30,  Ar-n).  Ventral  cu-ri  short, 
conical.  Proboscis  with  22  long  terminal  papillae  (10  pairs  bifid,  and 
a  single  dorsal  and  ventral  one),  with  22  longitudinal  rows  of  sub- 
terminal  papillae,  four  to  eight  per  row,  decreasing  in  size  basally. 

Key  to  the  species  of  Nephtys  from  Point  Barrow 

1.  Branchiae  cirriform,  sickle-shaped,  with  dorsal  'curi  digitiform  (fig.  30,  I,  n). 

Proboscis  with  a  long  median  dorsal  papilla 2 

Branchiae  foliaceous  or  cirriform  and  inflated  basally,  rudimentary  or  absent 
on  last  15-30  segments,  with  dorsal  cirri  flattened,  triangular,  pointed  (fig. 
30,  k,  m).  Proboscis  without  a  long  median  dorsal  papilla;  proximal  surface 
of  proboscis  ridged  (may  be  low  tubercles  in  N.  discors) 3 

2.  Branchiae  begin  on  setigers  3  or  4,  continuing  to  almost  posterior  end;  acicular 

lobes  rounded  or  sUghtly  bilobed,  especially  anteriorly  (fig.  30,  I).  Neuro- 
podial  postsetal  lamellae  extend  well  beyond  the  acicular  lobes,  bilobed  or 
irregularly  sinuous.  Setae  very  long,  flowing.  Proximal  surface  of  pro- 
boscis smooth  or  furrowed N.  longosetosa 

Branchiae  begin  on  setigers  5-8,  rudimentary  on  last  20-30  segments.  Acicular 
lobes  deeply  bilobed  in  anterior  and  median  regions  (fig.  30,  n) .  Neuropodial 
postsetal  lamellae  about  same  length  as  or  only  slightly  surpassing  the 
acicular  lobes.     Setae  shorter.     Proximal  surface  of  proboscis  tubercled. 

N.  ciliata 

3.  Branchiae  begin  on  setiger  6,  inflated  basally,  digitiform,  recurved  distally 

(fig.  30,  m).  Acicular  lobes  unequally  bilobed,  rounded  in  posterior  region. 
Postsetal  lamellae  of  anterior  and  median  regions  large,  foliaceous,  extending 
well  beyond  the  acicular  lobes;  in  posterior  region,  extend  only  slightly 

beyond N.  discors 

Branchiae  begin  on  setigers  8-14,  thick,  wide,  foliaceous  (fig.  30,  k).  Acicular 
lobes  rounded  to  conical.  Postsetal  lamellae  shorter  than,  as  long  as,  or 
only  slightly  longer  than  the  acicular  lobes N.  paradoxa 

261112—54 5 


268  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  los 

Nephtys  longosetosa  Oersted,  1843 

FlGUBE   30,  I 

Nephtys  longosetosa  Oersted,  1843,  p.  195,  pi.  6,  figs.  75,  76. — Hartman,  1944a, 
p.  339,  pi.  15,  fig.  7. 

Nephthys  longisetosa  not  Malmgren,  1865,  p.  106,  pi.  12,  fig.  20. — Eliason,  1920, 
p.  32. 

Nephthys  coeca  Murdoch,  1885,  p.  153  (part;  includes  var.  longisetosa;  not  Nereis 
caeca  Fabricius,  1780). 

Nephthys  longosetosa  Fauvel,  1923,  p.  367,  fig.  143,  f-h. — Monro,  1928,  p.  82. — 
Annenkova,  1937,  p.  163;  1938,  p.  162.— Friedrich,  1939,  p.  123.— Berkeley 
and  Berkeley,  1943,  p.  130;  1948,  p.  52,  fig.  76.— Gorbunov,  1946,  p.  38.— 
Wesenberg-Lund,  1950a,  p.  21;  1950b,  p.  60;  1951,  p.  44. 

Nephthys  ciliata  forma  longosetosa  Augener,  1928,  p.  698. 

Nephthys  ciliata  Treadwell,  1937,  p.  28  (part;  not  Nereis  ciliata  Muller,  1789). 

Description. — Length  up  to  170  mm.,  width  without  setae  6  mm. 
Tentacular  cirri  subequal  to  antennae.  Notopodial  postsetal  lamellae 
unequally  bilobed  (a  rather  large  upper  lobe  and  small  lobe  just  above 
the  dorsal  cirri),  extending  slightly  or  not  beyond  the  acicular  lobes. 
Neuropodial  postsetal  lamellae  of  anterior  region  rounded,  entire, 
extending  beyond  the  acicular  lobes;  those  of  the  median  and  posterior 
regions  large,  bilobed  or  irregularly  sinuous  on  ventral  side,  extending 
well  beyond  the  acicular  lobes.  Color:  In  alcohol:  Without  color, 
or  dusky,  especially  anteriorly. 

New  records. — Arctic  Alaska:  Eluitkak,  Elson  Lagoon  near  Point 
Barrow;  Point  Barrow  base,  washed  ashore;  off  Point  Barrow  base, 
1.7-25  fms.,  on  bottoms  of  gravel,  stones,  and  mud  (7  stations,  16 
specimens);  Cape  Lisburne,  5-7  fms.,  Dall.  Bering  Sea:  60°16'  N., 
167°41'  W.,  Stoney,  1884;  66°45'  N.,  166°35'  W.,  10  fms.,  Dall,  1880. 
Canadian  Arctic:  Duckett  Cove,  Hurd  Channel,  MelvUle  Peninsula, 
Foxe  Channel,  Bartlett,  1933.  East  Coast  North  America:  Off 
Labrador,  8  fms..  Blue  Dolphin  Expedition,  1949. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian,  Alaskan, 
and  Canadian  Arctic,  Greenland,  Spitsbergen,  Franz  Josef  Land, 
Barents  Sea,  White  Sea,  Novaya  Zemlya,  Kara  Sea.  Also  Iceland, 
Faroes  to  France;  to  Maine;  to  Lower  California  and  Panamd;  north 
Japan  Sea;  Strait  of  Magellan.    In  low  water  to  528  fathoms. 


Figure  30. — Nereidae:  a,  Nereis  pelagica,  dorsal  view  anterior  end;  b,  same,  middle  para- 
podium;  c.  Nereis  zonata,  middle  parapodium;  d,  compound  heterogomph  falciger;  e, 
compound  homogomph  falciger;  /,  compound  heterogomph  spiniger;  g,  compound  homo- 
gomph  spiniger;  h.  Nereis  zonata,  dorsal  view  extended  proboscis  showing  areas  of  parag- 
naths;  i,  same,  ventral  view.  Nephtyidae:  /,  Nephtys  paradoxa,  dorsal  view  anterior 
end;  k,  same,  middle  parapodium,  anterior  view;  /,  Nephtys  longosetosa,  middle  para- 
podium, anterior  view;  m,  Nephtys  discors,  middle  parapodium,  anterior  view;  n,  Nephtys 
ciliata,  middle  parapodium,  anterior  view.     (For  explanation  of  symbols,  see  p.  210.) 


MARINE    POLYCHAETE    WORMS — PETTIBONE 


269 


FiGtrRE  30. — For  explanation  see  facing  page. 


270  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  los 

Nephtys  ciliata  (Miiller,  1789) 

Figure  30,  n 

Nereis  ciliata  Miiller,  1789,  p.  14,  pi.  89,  figs.  1-4. 

Nephthys  ciliata  Malmgren,  1865,  p.  104,  pi.  12,  fig.  17. — Webster  and  Benedict, 

1887,  p.  709.— Moore,  1903,  p.  433.— Eliason,  1920,  p.  31.— Fauvel,  1923, 

p.  371,  fig.  145,  a-b;  1933,  p.  39.— Augener,  1928,  p.  699.— Gustafson,  1936, 

p.  7.— Treadwell,  1937,  p.  28  (part) .— Annenkova,  1937,  p.  164;  1938,  p. 

162.— Friedrich,  1939,  p.  123.— Gorbunov,  1946,  p.  38.— Thorson,  1946,  p. 

69,  figs.  32,  33.— Berkeley  and  Berkeley,  1948,  p.  55,  fig.  82.— Wesenberg- 

Lund,  1950a,  p.  21;  1950b,  p.  59;  1951,  p.  45. 
Nephthys  caeca  Murdoch,  1885,  p.  153  (part;  includes  var.  ciliata). — Moore,  1911, 

p.  243  (not  Nereis  caeca  Fabricius,  1780). 
Nepthys  ciliata  Chamberlin,  1920,  p.  9. 
Nepthys  hudsonica  Chamberlin,  1920,  p.  10,  pi.  2,  figs.  4-6. 
Nephtys  ciliata  Hartman,  1944a,  pp.  335,  339,  pi.  15,  fig.  9  (as  N.  incisa,  not  fig. 

10  {=N.  caeca));  1950,  p.  95. 

Description. — Length  100-300  mm.,  width  without  setae  9-11  mm. 
Tentacular  cirri  subequal  to  antennae.  Acicular  lobes  deeply  bilobed 
in  anterior  and  median  regions;  rounded  or  slightly  bilobed  in  posterior 
region.  Postsetal  lamellae  of  both  notopodia  and  neuropodia  rounded, 
entire,  short,  about  same  length  as  or  only  slightly  surpassing  the 
acicular  lobes.  Color:  In  alcohol:  White,  tannish,  or  anterior  third 
reddish  brown  or  buff.  (According  to  G.  E.  MacGinitie,  the  pellets 
of  this  worm  indicate  that  it  is  a  detritus  feeder.) 

New  records. — Arctic  Alaska:  Off  Point  Barrow  aero  radio  mast, 
10  fms.;  off  Point  Barrow  base,  up  to  8  miles  from  shore,  6-75.5  fms., 
on  bottoms  of  mud  and  various  combinations  of  mud,  pebbles,  gravel, 
stones,  rocks,  large  perforated  rocks,  and  shells  (18  stations,  37  speci- 
mens). Siberia:  Plover  Bay,  Dall,  1880.  Alaska:  Chiachi  Island,  20 
fms.,  mud;  Round  Island,  Coal  Harbor,  6-8  fms.,  mud,  1872;  Port 
Mulgrove,  Yakutat  Bay,  6-40  fms.,  1874;  Port  Etches,  12-18  fms., 
1874;  aU  collected  by  DaU.  West  Coast  North  America:  Wash- 
ington and  Puget  Sounds,  low  water  to  83  fms.  (common),  Petti- 
bone.  East  Coast  North  America:  Off  Labrador,  6-95  fms..  Blue 
Dolphin  Expeditions,  1949,  1950,  1951;  off  Nova  Scotia,  Bay  of 
Fundy,  Maine,  Massachusetts,  14-110  fms.,  U.  S.  Fish  Commission. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian,  Alaskan, 
aud  Canadian  Arctic,  Greenland,  Spitsbergen,  Norway,  Barents  Sea, 
White  Sea,  Novaya  Zemlya,  Kara  Sea.  Also  Iceland,  Faroes  to 
France;  to  Massachusetts;  to  Southern  California;  to  Japan,  China. 
In  low  water  to  500  fathoms. 

Nephtys  discors  Ehlers,  1868 

Figure  30,  m 

Nephthys  discors  Ehlers,  1868,  p.  626,  pi.  23,  figs.  39,  40.— Verrill,  1873,  p.  103.— 

Webster  and  Benedict,  1887,  p.  709. 
Nephtys  discors  Hartman,  1938c,  p.  9,  pi.  1,  figs.  2,  3;  1950,  p.  96. 
Nephtys  rickettsi  Hartman,  1938b,  p.  153;  1950,  p.',97. 


MARINE    POLYCHAETE    WORMS — PETTIBONE  271 

Description. — Length  115-300  mm.,  width  without  setae  9-12  mm. 
Dorsal  pair  tentacular  cuTi  about  half  as  large  as  ventral  pair. 
Branchiae  begin  on  setiger  6,  at  first  a  small  swelling  on  the  lower  side 
of  the  base  of  the  flattened,  triangular  dorsal  cirrus,  gradually  becom- 
ing larger;  where  best  developed,  they  are  inflated  on  the  basal  half, 
with  the  distal  half  more  slender,  digitiform  and  slightly  recurved  or 
hooked;  they  are  smaller  and  rudimentary  only  on  posterior  half  of 
body.  Notopodial  acicular  lobes  of  anterior  and  middle  region 
unequally  bilobed,  the  upper  one  wider  and  shorter  and  the  ventral 
one  narrower  and  longer;  rounded  in  posterior  region.  Neuropodial 
acicular  lobes  slightly  bilobed  in  anterior  region;  very  low  and  rounded 
in  middle  and  posterior  regions  (not  very  distinct  from  posterior 
lamellae).     Color:  In  alcohol:  Brownish. 

Remarks. — The  type  of  N.  rickettsi  Hartman  from  Alaska  was 
compared  with  a  specimen  of  A^.  discors  from  Bay  of  Fundy  and  is 
herein  referred  to  the  latter  species. 

New  records. — Arctic  Alaska:  Point  Barrow  base,  washed  ashore; 
off  Point  Barrow  base,  up  to  12.1  miles  from  shore,  24.7-123.5  fms., 
on  bottoms  of  mud,  gravel-mud,  stones  and  large  perforated  rocks, 
and  mass  of  worm  tubes  (5  stations,  5  specimens).  West  Coast 
North  America:  Puget  and  Washington  Sounds,  70-83  fms.,  mud 
(3  specimens),  Pettibone,  1938,  1939.  East  Coast  North  America: 
Bay  of  Fundy,  U.  S.  Fish  Commission. 

Distribution. — Alaskan  Arctic  to  southern  California;  Bay  of  Fundy 
to  Maine.     In  24-268  fathoms. 

Nephtys  paradoxa  Malm,  1874 

Figure  30,  j,  k 

Nephthys  paradoxa  Malm,  1874,  p.  78,  pi.  1,  fig.  2. — Fauvel,  1923,  p.  375,  fig. 
146,  f-i.— Augener,  1928,  p.  701.— Gustafson,  1936,  p.  7.— Annenkova,  1938, 
p.  163.— Friedrich,  1939,  p.  123.— Berkeley  and  Berkeley,  1943,  p.  130.— 
Wesenberg-Lund,  1950a,  p.  22;  1950b,  p.  61;  1951,  p.  47. 

Nephthys  phyllobranchia  Mcintosh,  1885,  p.  164,  pi.  26,  fig.  10;  pi.  27,  fig.  3;  pi. 
14,  A,  figs.  12,  13. 

?  Nephthys  brachycephala  Moore,  1903,  p.  431. — Annenkova,  1937,  p.  164;  1938, 
p.  162.— Zatsepin,  1948,  p.  121,  pi.  30,  fig.  9. 

Nephtys  paradoxa  Hartman,  1944a,  pp.  335,  339,  pi.  15,  fig.  6;  1950,  p.  111. 

Nephtys  phyllobranchia  Hartman,  1950,  p.  111. 

Description. — Length  150-200  mm.,  width  without  setae  8-13  mm. 
Dorsal  pair  tentacular  cirri  reduced  to  a  tubercle,  smaller  than  ventral 
pair.  Branchiae  begin  on  setigers  11-13  (8-14),  very  short,  triangular 
at  first,  gradually  becoming  larger;  where  best  developed,  they  are 
flat,  wide  foliaceous,  weakly  to  distinctly  concave;  they  are  small, 
nearly  absent  on  the  last  15-30  segments.  The  anterior  acicular 
lobes  are  rounded;  those  of  the  middle  and  posterior  regions  are 
conical,  pointed.     The  postsetal  lamellae  in  the  anterior  region  are 


272  PROCEEDINGS    OF   THE   NATIONAL   MUSEUM  vol.  103 

slightly  longer  than  the  acicular  lobes;  they  are  shorter  than  the 
conical  acicular  lobes  in  the  middle  and  posterior  regions.  Color: 
In  alcohol :  Dusky,  especially  anteriorly. 

Remarks. — Augener  (1928)  indicated  that  he  was  of  the  opinion 
that  N.  phyllobranchia  Mcintosh  should  be  referred  to  A^,  paradoxa; 
Hartman  (1950)  stated  that  they  are  separable  by  only  slight  differ- 
ences. The  type  of  A^.  brachycephala  Moore  from  Japan  is  unsatis- 
factory (dried,  when  received);  the  description  is  in  agreement  with 
N.  paradoxa. 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  up  to  12.1 
miles  from  shore,  23.5-123.5  fms.,  on  bottoms  of  mud  and  mass  of 
worm  tubes  (2  stations,  3  specimens). 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian,  Alaskan, 
and  Canadian  Arctic,  Greenland,  Spitsbergen,  Norway,  Barents  Sea, 
Kara  Sea.  Also  Iceland,  Faroes  to  France;  off  Delaware;  Japan. 
In  3.3-4,001  fathoms. 

Family  Glyceridae 

Body  rounded,  elongated,  attenuated  at  both  ends,  not  divided 
into  two  regions,  either  only  uniramous  or  only  biramous.  Segments 
bi-  or  tri-annulate.  Prostomium  sharply  conical,  wide  basaUy,  with 
a  pair  of  nuchal  organs,  transversely  annulated,  ending  in  four  minute 
antennae  distally  (fig.  31,  a).  Small,  globular  dorsal  cirri;  larger 
ventral  cirri  (fig.  31,  b,  c);  two  anal  cirri.  Branchiae  present  or 
absent.  Notosetae  simple  capillaries;  neurosetae  compound  spinigers. 
Proboscis  eversible,  long,  clavate,  with  numerous,  small  papillae 
(proboscidial  organs)  and  four  dark,  falcate,  horny  jaws  each  with  an 
attached,  embedded,  rodlike  aileron  (fig.  31,  d).  Epitoky  affects 
entke  individual  (parapodial  lobes  become  more  elongate,  setae  longer 
and  more  numerous). 

Represented  by  a  single  species  from  Point  Barrow. 

Genus  Glycera  Savigny,  1818 
Glycera  capitata  Oersted,  1843 

FiGUBE  31,  a-d 

Glycera  capitata  Oersted,  1843,  p.  196,  pi.  7,  figs.  87,  88,  90-94,  96,  99.— Ehlers, 
1913,  p.  603.— Fauvel,  1923,  p.  385,  fig.  151,  a-e.— Augener,  1928,  p.  734.— 
Monro,  1930,  p.  115;  1936,  p.  114.— Annenkova,  1937,  p.  165;  1938,  p.  164.— 
St0p-Bowitz,  1941,  p.  186,  fig.  1;  1948b,  p.  4.— Hartman,  1944a,  pp.  336, 
339;  1948,  p.  28;  1950,  p.  76,  pi.  11,  figs.  1-4.— Gorbunov,  1946,  p.  38.— 
Zatsepin,  1948,  p.  117,  pi.  31,  fig.  9.— Berkeley  and  Berkeley,  1948,  p.  38, 
fig.  52. — Hartman  and  Reish,  1950,  p.  20. — Wesenberg-Lund,  1950a,  p.  23; 
1950b,  p.  64;  1951,  p.  49. 

Glycera  setosa  Oersted,  1843,  p.  198,  pi.  7,  figs.  89,  95,  97  (epitokous  form). 


MARINE    POLYCHAETE    WORMS — PETTIBONE  273 

Glycera  nana  Johnson,  1901,  p.  411,  pi.  10,  fig.  103.— Berkeley  and  Berkeley, 

1948,  p.  37,  figs.  50,  51. 
Glycera  lapidum  Fauvel,  1914,  p.  205;  1923,  p.  386,  fig.  151, /-m;  1934a,  p.  41.— 

Eliason,   1920,  p.  36,  fig.  6.— Treadwell,   1937,  p.  32.— St0p-Bowitz,  1941, 

p.  191,  fig.  2;  1948b,  p.  7. 

Description. — Length  80  mm.,  width  without  setae  6  mm.  (length 
up  to  150  mm. — Hartman,  1950).  Segments  usually  triannulate 
(may  be  biannulate,  a  slightly  longer  anterior  ring  at  the  level  of  the 
parapodia  and  a  shorter  posterior  ring;  the  former  may  be  sub- 
divided, thus  triannulate;  the  latter  may  also  be  somewhat  divided, 
thus  tetra-annulate) .  Conical  prostomium  with  8-16  annuli  (often 
difficult  to  count  accurately).  Dorsal  cirri  small,  globular,  inserted 
well  above  the  parapodia.  Ventral  cirri  wide,  conical.  Parapodia 
biramous  throughout;  notopodia  represented  by  a  short,  conical 
presetal  lobe  and  a  small  bundle  of  simple  capillary  setae;  neuropodia 
with  a  longer,  conical  presetal  lobe  and  a  fan-shaped  group  of  com- 
pound spinigerous  setae;  a  single  short,  wide,  rounded  postsetal  lobe 
common  to  both  notopodia  and  neuropodia  (thus  two  conical  an- 
terior lobes  and  a  single  postsetal  lobe).  The  relative  length  of  the 
parapodial  lobes  may  be  variable,  associated  with  the  epitokous 
condition,  as  indicated  by  Hartman  (1950).  Proboscis  with  two 
kinds  of  papillae,  numerous,  long,  cylindrical,  slender  ones,  and 
fewer,  larger,  short,  subspherical  ones;  aileron  of  jaws  with  a  long 
lateral  prolongation.     Color:  In  alcohol:  Brownish. 

Remarks. — Monro  (1930)  referred  Qlycera  lapidum  Quatrefages  to 
G.  capitata;  they  have  been  separated  on  minor  and  apparently 
variable  characters;  Fauvel  (1914,  1923)  regarded  them  as  mere 
varieties.     Hartman  (1950)  referred  G.  nana  Johnson  to  G.  capitata. 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  15  miles 
from  shore,  78.2  fms.,  on  bottom  of  small  rocks  with  worm  tubes  (1 
station,  1  specimen).  Northwest  Greenland:  Off  Conical  Rock, 
20-40  fms.,  1938;  1  mile  northwest  Conical  Rock,  1940;  west  side 
Wolstemholm  Island,  1940;  all  collected  by  Bartlett.  East  Coast 
North  America:  Off  Nova  Scotia,  Maine,  Massachusetts.  Rhode 
Island,  low  water  to  410  fms.,  U.  S.  Fish  Commission.  Alaska: 
Eastern  Harbor,  Sitka,  gravelly-mud,  15  fms.,  1874;  between  Pin- 
nacle and  Ulakhta,  Unalaska,  16  fms.,  1874;  Captain's  Harbor, 
Unalaska,  25-75  fms.,  coarse  sand,  1874;  all  collected  by  Dall. 
Albatross  Sta.  Herendeen  Bay,  1890.  Washington:  Albatross  Sta. 
2876,  48°33'  N.,  124°53'  W.,  59  fms.,  and  Sta.  2879,  48°53'  N.,  125°53' 
W.,  34  fms.,  1888;  Strait  of  Juan  de  Fuca,  Washington  and  Puget 
Sounds,  low  water  to  165  fms.,  Pettibone. 

Distribution. — Widely  distributed  in  the  Arctic :  Siberian,  Alaskan, 
and  Canadian  Arctic,  Baffin  Bay,  Davis  Strait,  Greenland,  Spits- 
bergen, Norway,  Barents  Sea,  White  Sea,  Novaya  Zemlya,  Kara 


274  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  m 

Sea.  Also  Iceland,  Faroes  to  Portugal,  Azores,  Madeira,  Mediter- 
ranean, Adriatic;  to  Rhode  Island;  to  Gulf  of  California  and  Mexico; 
Japan;  South  Atlantic  and  Antarctic.  In  low  water  to  1,889  fathoms; 
epitokous  phase  in  surface  layers. 

Family  Goniadidae 

Body  elongated,  more  or  less  attenuated  at  both  ends,  divided 
into  two  regions,  anterior  region  with  uniramous  parapodia  (fig.  31, 
f)  and  long,  wider  posterior  region  with  biramous  parapodia  (fig.  31, 
g).  Segments  uniannulate.  Prostomium  sharply  conical,  trans- 
versely annulated,  ending  distally  in  four  small  antennae  (fig.  31,  e). 
Elongated  conical  dorsal  and  ventral  cirri  (fig.  31,  /,  g).  Two  anal 
cirri.  Notosetae  simple;  neurosetae  compound.  Branchiae  absent. 
Proboscis  eversible,  long,  cylindrical,  with  terminal  papillae  and 
proboscidial  organs,  a  pair  of  larger,  dentate,  dark  horny  macrognaths, 
and  a  circlet  of  many  smaller  H-  or  Y-shaped  micrognaths.  Epitoky 
involves  only  biramous  portion. 

Represented  by  a  single  species  from  Point  Barrow. 

Genus  Glycinde  F.  Miiller,  1858 
Glycinde  wireni  Arwidsson,  1898 

FiGUEE  31,  e-g 

Goniada  nordmanni  Wir^n,  1883,  p.  403,  pi.  30,  figs.  4,  5,  pi.  32,  figs.  1,  2;  (not 

Eone  nordmanni  Malmgren,  1867). 
Glycinde  wirini  Arwidsson,  1898,  p.  53,  pi.  3,  figs.  48,  49. — not  Moore,  1908,  p. 

348  (  =  (?.  armigera  Moore;  in  USNM). 

Description. — Length  up  to  45  mm.,  width  without  setae  2.5  mm., 
width  including  setae  5  mm.,  120  segments.  Anterior  region  rounded, 
tapering  sharply  to  pointed  anterior  end;  posterior  two-thirds  wider, 
more  flattened,  tapering  gradually  posteriorly.  Prostomium  with  10 
annuli;  two  pairs  of  small  deeply-set  eyes,  one  pair  in  basal  ring,  one 
pair  in  distal  ring  (may  be  obscure).  Anterior  uniramous  region 
consisting  of  30-32  (31  in  Arwidsson)  setigers,  with  presetal  lobe  wide 
basally,  narrowed  abruptly  to  a  lanceolate  tip;  postsetal  lobe  wide, 
conical;  pre-  and  postsetal  lobes  as  well  as  dorsal  and  ventral  cirri 
subequal  in  length;  a  fan-shaped  bundle  of  compound,  spinigerous 
neurosetae.  Posterior  biramous  region  with  shorter  notopodia  dis- 
tinctly separated  from  neuropodia.  Notopodium  a  short,  rounded 
lobe  with  a  short  dorsal  cirrus  and  a  short,  protruding  presetal  lobe; 
notosetae  few  in  number,  acicular,  hooked  at  tip,  with  long  pointed 
hood.  Neuropodia  with  short  conical,  postsetal  lobe,  subequal  to 
ventral  cirrus;  presetal  lobe  wide  basally  with  a  lanceolate  narrower 
tip,  longer  than  postsetal  lobe;  neurosetae  as  in  anterior  region. 


MARINE    POLYCHAETE    WORMS — PETTIBONE  275 

except  they  may  be  longer  and  more  numerous.  Proboscis  without 
chevrons;  proboscidial  organs  (horny,  yellow,  spinous  processes)  of 
several  kinds  arranged  in  longitudinal  series.  Color:  In  alcohol: 
Uniformly  greenish  brown  or  somewhat  banded  with  one  wide,  dark- 
greenish-brown  band  per  segment;  iridescent. 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  up  to  4 
miles  from  shore,  23  to  29.1  fms.,  on  various  combinations  of  mud, 
sand,  gravel,  stones,  rocks,  shells  (3  stations,  5  specimens). 

Distribviion. — Arctic  Alaska  to  Bering  Sea.     In  23  to  29  fathoms. 

Family  Lumbrineridae 

Body  elongate,  cylindrical,  tapered  slightly  anteriorly  and  more 
strongly  posteriorly,  superficially  resembling  oligochaetes.  Pro- 
stomium  conical  to  rounded,  with  a  pair  of  nuchal  organs  on  posterior 
margin,  without  antennae  or  palps  (fig.  31,  h).  First  two  segments 
apodous  and  achaetous,  without  tentacular  cirri.  Without  dorsal 
and  ventral  cirri.  Two  to  four  anal  cirri.  Parapodia  imiramous. 
Proboscis  eversible,  with  dark,  chitinous,  well-developed  masticatory 
apparatus  consisting  of  a  pair  of  ventral  mandibles  and  four  pairs  of 
maxillary  plates  (fig.  31,  i,j). 

Represented  by  a  single  species  from  Point  Barrow. 

G^nus  Lumbrineris  Blainville,  1828 
Lumbrineris  fragilis  (O.  F.  Miiller,  1776) 

Figure  31,  h-n 

Lumbricus  fragilis  Miiller,  1776,  p.  216;  1788,  p.  22,  pi.  22,  figs.  1-3. 

Lumbrinereis  fragilis  Malmgren,  1867,  pi.  14,  fig.  83. — Chamberlin,  1920,  p.  15. — 
Eliason,  1920,  p.  33.— Berkeley  and  Berkeley,  1943,  p.  130. 

Lumbriconereis  minuta  Th^el,  1879,  p.  42,  pi.  4,  figs.  57-59. — Fauvel,  1911,  p. 
22.— Augener,  1928,  p.  732.— Wesenberg-Lund,  1950a,  p.  28;  1950b,  p.  71. 

Lumbriconereis  fragilis  Fauvel,  1923,  p.  430,  fig.  171,  k,  I. — Augener,  1928,  p. 
730,  pi.  11,  fig.  5.— Annenkova,  1937,  p.  167;  1938,  p.  167.— Gorbunov,  1946, 
p.  39.— Thorson,  1946,  p.  74.— Zatsepin,  1948,  p.  124,  pi.  31,  fig.  14,— 
Wesenberg-Lund,  1950a,  p.  27;  1950b,  p.  69;  1951,  p.  55. 

Lumbrineris  fragilis  Hartman,  1944a,  pp.  335,  340. 

Description. — Length  150-250  mm.,  width  without  setae  up  to  7 
mm.,  width  with  setae  12  mm.  Prostomium  conical,  pointed.  First 
two  achaetous  segments  about  equal  to  the  following  segments  or  the 
first  one  slightly  longer.  Parapodia  without  branchiae,  with  presetal 
lobe  short,  rounded ;  postsetal  lobe  in  anterior  region  of  body  slightly 
longer,  rounded,  diagonally  truncate;  in  middle  and  posterior  regions 
of  body,  postsetal  lobe  extended  somewhat  dorsally,  somewhat  digiti- 
form;  with  arched  limbate  setae  with  fine  capillary  tips,  dark  amber- 
colored  on  basal  half,  on  first  60-100  setigers;  with  simple  (not  com- 
pound), hooded,  hooked  setae,  dark  amber-colored  basally,  beginning 


276  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  m 

on  setigers  15-24  (22-35  or  more) ;  on  young  specimens,  the  hooded, 
hooked  setae  may  begin  on  more  anterior  segments;  acicula  black. 
With  two  anal  cu'ri  on  ventral  side  of  short,  disclike  pygidium.  Jaw 
formula  of  proboscis  (numbers  refer  to  number  of  teeth) :  M  II,  4+4 
(sometimes  5?);  M  III,  1  +  1  (or  2  +  1);  M  IV,  1  +  1.  Color:  In  hfe: 
Reddish  orange,  iridescent.  In  alcohol:  Colorless  to  gray,  iridescent; 
young  specimens  uniformly  reddish  brown  or  somewhat  banded  with 
one  wide  dark  brown  band  per  segment. 

Remarks. — Three  small  specimens  (0.8  mm.  wide  without  setae) 
have  the  hooded,  hooked  setae  beginning  on  the  first  one  to  three 
setigers  as  in  L.  minuta  Theel.  They  are  considered  to  be  the  young 
of  L.  jragilis  as  has  been  suggested  earlier  by  Levinsen  (1882)  and 
Eliason  (1920). 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  up  to  12.1 
miles  from  shore,  20-123.5  fms.,  on  bottoms  of  mud,  stones,  worm  tubes 
and  various  combinations  of  mud,  sand,  gravel,  stones,  rocks,  shells 
(16  stations,  29  specimens).  East  Greenland:  Off  Cape  Hold  with 
Hope,  23-40  fms.,  Bartlett,  1939.  Eastern  North  America:  Off 
Labrador,  5-13  fms..  Blue  Dolphin  Expeditions,  1949,  1950;  off  Nova 
Scotia,  Bay  of  Fundy,  Maine,  Massachusetts,  Rhode  Island,  7-452 
fms.,  U.  S.  Fish  Commission;  Woods  Hole  region,  low  water  and 
dredged,  Pettibone,  1950,  1951.  Alaska:  Chiachi  Islands,  20  fms., 
mud,  Dall,  1874. 

Distrih'ution. — Widely  distributed  in  the  Arctic:  Siberian  and  Alas- 
kan Arctic,  Davis  Strait,  Greenland,  Jan  Mayen,  Spitsbergen,  Franz 
Josef  Land,  Barents  Sea,  Novaya  Zemlya,  Kara  Sea.  Also  Iceland, 
Faroes,  Norway  to  Azores,  Madeira,  Mediterranean;  Hudson  Bay  to 
Rhode  Island;  Bering  Sea;  north  Japan  Sea.  In  low  water  to  1,883 
fathoms. 

?  Larval  Lumbrineris  fragilis  (O.  F.  Miiller) 

On  two  occasions  nonpelagic  larval  polychaetes  were  found  in  a 
mucous  mass  off  Point  Barrow  base:  September  9,  1948,  21  fathoms, 
from  breaking  rock  (the  larvae  described  as  rose-red);  September  6, 
1949,  36  fathoms,  attached  to  tunicate  Boltenia  echinata.    Prostomium 


Figure  31. — Glyceridae:  a,  Glycera  capiiata,  dorsal  view  prostomium;  b,  same,  parapodium 
from  anterior  region,  posterior  view;  c,  same,  parapodium  from  middle  region,  anterior 
view;  d,  same,  one  of  jaws.  Goniadidae:  e,  Glycinde  tvireni,  dorsal  view  anterior  end; 
/,  same,  uniramous  parapodium  from  anterior  region,  anterior  view;  g,  same,  biramous 
parapodium  from  posterior  region,  anterior  view.  Lumbrineridae:  A,  Lumbrineris 
fragilis,  dorsal  view  anterior  end;  i,  same,  maxillary  plates  of  jaw  apparatus  (one  of  each 
pair  shown);  j,  same,  mandibles  of  jaw  apparatus;  k,  same,  parapodium  from  anterior 
region,  posterior  view;  /,  same,  parapodium  from  posterior  region,  posterior  view;  m, 
same,  limbate  seta  with  capillary  tips;  n,  same,  simple  hooded  hooked  seta  (crotchet). 
(For  explanation  of  symbols  see  p.  210.) 


MARINE    POLYCHAETE    WORMS — PETTIBONE 

Ir.  an  -dC       "PO^, 


277 


J.        ^  -  N 

Figure  31. — For  explanation  see  facing  page. 


278  PROCEEDINGS   OP   THE   NATIONAL  MUSEUM  vol.  i03 

conical,  without  appendages.  Two  wide  peristomial  segments,  the 
second  with  wide  transverse  ciliated  band.  Setigerous  segments  8, 
gradually  tapering  posteriorly,  each  parapodium  with  single  limbate 
capillary  seta  and  hooded  seta.  Pygidium  bulbous,  with  wide  trans- 
verse ciliated  band. 

Family  Orbiniidae  (Ariciidae) 

Body  long,  vermiform,  with  very  numerous  segments,  divided  into 
two  weakly  to  sharply  separable  regions:  (1)  thoracic:  More  or  less 
flattened  dorsoventrally  and  enlarged;  attenuated  anteriorly;  neuro- 
podia  cushionlike,  with  several  rows  of  setae;  notopodia  more  or  less 
cirriform,  a  bundle  of  crenulate  capillary  setae  (fig.  32,  6);  (2)  ab- 
dominal: Much  longer,  semicylindrical;  neuropodia  bilobed,  project- 
ing dorsally  (fig.  32,  c).  Branchiae  medial  to  notopodia,  simple, 
dorsal,  erect,  lanceolate  or  straplike,  strongly  ciliated,  a  pair  on  aU 
segments  except  the  anterior  ones.  Prostomium  conical  or  globular, 
without  appendages,  with  or  without  two  small  eyes,  with  two  ciliated 
nuchal  organs  (fig.  32,  a).  First  one  or  two  segments  apodous  and 
achaetous.  Usually  without  dorsal  or  ventral  cirri.  Setae  simple,  of 
varied  form.  Pygidium  with  two  to  four  anal  cirri.  Proboscis 
eversible,  unarmed,  a  voluminous,  weakly-  to  much-branched,  soft  sac. 

Represented  by  a  single  species  from  Point  Barrow. 

Genus  Scoloplos  Blainville,  1828 
Scoloplos  armiger  (O.  F.  Miiller,  1776) 

Figure  32,  a-e 

Lumhricus  armiger  O.  F.  MuUer,  1776,  p.  215;  1788,  p.  22,  pi.  22,  figs.  4,  5. 

Aricia  arctica  Hansen,  1882,  p.  34,  pi.  5,  figs.  20-26. — Murdoch,  1885,  p.  154. 

Scoloplos  armiger  Fauvel,  1914,  p.  224;  1927,  p.  20,  fig.  6,  k-g. — Eliason,  1920, 
p.  39. — Augener,  1928,  p.  742.— Monro,  1930,  p.  145. — Annenkova,  1931,  p. 
203;  1937,  p.  169;  1938,  p.  170.— Okuda,  1937a,  p.  102,  fig.  4.— Wesenberg- 
Lund,  1939b,  p.  12;  1950a,  p.  29;  1950b,  p.  73;  1951,  p.  58.— Berkeley  and 
Berkeley,  1942,  p.  195;  1952,  p.  97,  figs.  197-199.— Hartman,  1944a,  pp.  336, 
340,  pi.  18,  fig.  5  (not  pi.  19,  fig.  6  (=Orbinia)) .—Thorson,  1946,  p.  78,  fig. 
37.— Gorbunov,  1946,  p.  39.— Zatsepin,  1948,  p.  128,  pi.  32,  fig.  1. 

Haploscoloplos  alaskensis  Hartman,  1948,  p.  30,  fig.  8,  a-c. 

Description. — ^Length  up  to  120  mm.,  width  2  mm.  Prostomium 
conical,  pointed,  with  two  deeply  buried  eyes  (not  visible  when  pre- 
served). First  segment  achaetous  and  apodous.  Branchiae  first 
appear  on  setigers  12  or  13  (9-17),  very  small  at  first,  then  triangular, 
then  ligulate.  Thoracic  region  consisting  of  about  17  setigers 
(12-20) ;  notopodia  with  conical  postsetal  lobes  and  numerous,  finely 
crenulate  setae  with  capillary  tips;  neuropodia  elongate,  cushionlike, 
with  a  median  postsetal  conical  lobe,  with  numerous  rows  of  crenulate 


MARINE    POLYCHAETE    WORMS — PETTIBONE  279 

setae  with  capillary  tips;  some  of  these  setae  in  the  lower  part  of  the 
neuropodial  bundle  may  be  worn  down,  with  blunt  tips  resembling 
crotchets.  These  so-called  crotchets  may  be  more  or  less  numerous 
on  all  thoracic  segments  or  on  some  only,  or  may  be  lacking.  Ab- 
dominal region  having  an  exceedingly  straggly  appearance  with  the 
dorsally  directed  branchiae  and  parapodial  lobes;  notopodia  with 
erect,  digitiform  postsetal  lobes  and  capillary  crenulate  setae  and 
sometimes  also  with  forked  setae;  neuropodia  with  two  conical, 
unequal  lips,  with  capillary  crenulate  setae  and  an  elongate  thickened 
parapodial  flange  below  the  neuropodial  lobe.  On  the  last  few  thoracic 
and  first  few  abdominal  segments  there  may  be  one  or  two  extra 
conical  podal  lobes,  one  at  the  level  of  the  lower  part  of  the  neuropodial 
setae  and  one  subpodal;  they  are  in  the  transitional  region  of  thoracic 
and  abdominal  regions  and  become  part  of  the  parapodial  flanges  of 
the  abdominal  segments.  Without  intermediate  interramal  cirri, 
but  with  lateral,  ciliated  organs  between  the  rami.  Pygidium  with 
two  filiform  anal  cirri.  Proboscis  soft,  more  or  less  lobed.  Color: 
In  alcohol:  Without  color  or  yellowish  brown,  with  darker  areas  on 
the  elongate  neuropodial  flanges  of  the  abdominal  region  and  on  the 
branchiae. 

Remarks. — Haploscoloplos  alaskensis  Hartman  from  southwestern 
Alaska  is  herein  referred  to  S.  armiger.  It  differs  from  the  typical 
S.  armiger  in  lacking  crotchets  in  the  thoracic  neuropodia.  As 
indicated  previously  by  Fauvel  (1914)  and  others,  this  seems  to  be  a 
variable  character,  perhaps  associated  with  the  substratum  and  age. 
The  crotchets  were  not  present  on  small  specimens  from  Point  Barrow, 
and  they  may  be  more  numerous  in  some  parapodia  than  in  others 
and  more  abundant  in  the  lower  part  of  the  neuropodia;  they  appear 
to  be  worn  crenulate  capillary  setae.  On  certain  substrata,  pre- 
sumably, the  setae  may  not  be  worn  and  thus  the  so-called  crotchets 
are  absent. 

New  records. — Arctic  Alaska:  Eluitkak  Pass,  Elson  Lagoon  near 
Point  Barrow;  west  side  Elson  Lagoon  near  entrance  to  small  lagoon 
to  the  west,  1.2  fms.;  off  Point  Barrow  base,  up  to  12.1  mQes  from 
shore,  1.7-75.5  fms.,  on  bottoms  of  mud,  stones,  and  various  com- 
binations of  mud,  gravel,  stones,  rocks,  large  perforated  rocks  (12 
stations,  19  specimens).  Between  Icy  Cape  and  Cape  Lisbume, 
15-20  fms.,  mud,  sand,  Dall,  1874.  East  Coast  North  America: 
Bay  of  Fundy,  Maine,  Massachusetts,  13—48  fms.,  U.  S.  Fish 
Commission. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian  and 
Alaskan  Arctic,  Davis  Strait,  Greenland,  Jan  Mayen,  Spitsbergen, 
Franz  Josef  Land,  White  Sea,  Novaya  Zemlya,  Kara  Sea.  Also 
Iceland,  Faroes,  Shetlands  to  France,  Mediterranean;  Labrador  to 


280  PROCEEDINGS   OF   THE   NATIONAL  MUSEUM  vol.  103 

Massachusetts;  southern  Alaska  to  British  Columbia;  Japan;  South 
Pacific  (Chile);  Antarctic;  off  South  Africa.  In  low  water  to  1,100 
fathoms. 

Family  Spionidae 

Body  vermiform,  without  distinct  regions.  Prostomium  small, 
elongated,  scarcely  separated  from  an  enlarged  peristomium  lateral  to 
prostomium,  with  or  without  eyes  (usually  four),  antennae  usually 
lacking  (may  be  a  small  occipital  antenna),  two  very  long,  extensile, 
tentaculiform,  longitudinally  grooved,  easily  deciduous  palps  (fig. 
32,  /,  i,  I,  r) .  Parapodia  biramous,  with  dorsal  and  ventral  postsetal 
lamellae  more  or  less  developed;  dorsal  and  ventral  cirri  lacking 
(fig.  32,  g,  s).  With  ligulate  or  cirriform  dorsal  branchiae  on  upper 
part  of  parapodial  bases  on  a  certain  number  of  segments  (rarely 
pinnate,  rarely  lacking).  Setae  simple  capillaries  and  hooded 
crotchets  (fig.  32,  h,  k,  q).  Pygidium  terminated  in  an  anal  cup 
(fig.  32,  p)  or  anal  cirri.  Proboscis  short,  scarcely  or  somewhat 
protrusible,  may  be  somewhat  lobulated. 

Represented  by  four  genera  and  four  species  from  Point  Barrow. 
All  genera  have  branchiae  present  on  certain  segments  and  prostomia 
without  frontal  horns. 

Key  to  the  genera  of  Spionidae  from  Point  Barrow 

1.  Setigerous   segment   5   strongly   modified   with   special   large   amber-colored 

setae  and  without  dorsal  and  ventral  lamellae  (fig.  32,  m,  n).  Branchiae 
numerous,  paired,  dorsal,  simple,  straplike.  Notopodia  with  capillary 
setae  only;  neuropodia  with  capillary  setae  and  hooded  bidentate  crotchets 
beginning  on  setigers  7  or  8  (fig.  32,  q).     Pygidium  ending  in  anal  cup 

(fig.  32,  p) Polydora  (p.  280) 

Setigerous  segment  5  not  modified 2 

2.  Branchiae  on    only  few  anterior   segments,  3-11   pairs,  often  pinnate   (fig. 

32,  j),  beginning  on  setiger  2.     Capillary  setae  and  hooded  crotchets  in  both 

noto-  and  neuropodia.    Pygidium  with  anal  cirri Prionospio  (p.  282) 

Branchiae  more  numerous,  simple,  straplike.     Notopodia  with  capillary  setae 
only;  neuropodia  with  capillary  setae  and  hooded  crotchets 3 

3.  Branchiae  begin  on  setiger  1,  continuing  up  to  last  setigers.     Pygidium  with 

anal  cirri Spio  (p.  284) 

Branchiae  begin  on  setiger  2,  continuing  on  large  number  of  segments.     Py- 
gidium with  anal  cup,  may  be  somewhat  lobulated Nerinides  (p.  285) 

Genus  Polydora  Rose,  1801 
Polydora  caulleryi  Mesril,  1897 

Figure  32,  l-q 

Polydora  caulleryi  Mesnil,  1897,  p.  88,  pi.  3,  figs.  12-16.— Fauvel,  1927,  p.  54, 

fig.  19,  f-h. 
Polydora  brachycephala  Hartman,  1936c,  p.  48,  figs.  3-5;  1944b,  p.  258. 


MARINE    POLYCHAETE    WORMS — PETTIBONE  281 

Description. — Up  to  50  mm.  long,  2.2  mm.  wide.  Prostomium 
feebly  notched  anteriorly  or  distinctly  bifid,  prolonged  posteriorly  in 
a  crest  extending  to  setiger  4  (to  6) ,  without  distinct  occipital  antenna, 
with  fom-  eyes  in  trapezoidal  arrangement  (or  six  in  young  specimens) 
or  eyes  lacking.  Palps  usually  missing  in  alcohol.  A  protrusible, 
somewhat  lobulated,  brownish  proboscidial  region.  Branchiae  begin 
on  setigers  7  or  8,  continuing  on  numerous  segments,  absent  from 
more  posterior  segments;  branchiae  short  (about  length  of  capillary 
notosetae),  straplike,  continuous  with  dorsal  lamellae.  Modified  seti- 
ger 5  with  group  of  capillary  notosetae,  usually  5-7  (3-12)  special 
amber-colored  hooks,  and  group  of  capillary  neurosetae;  special  hooks 
all  similar,  falcate,  with  pectinate  top.  With  conspicuous,  somewhat 
conical  dorsal  and.  ventral  postsetal  lamellae  on  setigers  1-4,  lacking 
on  modified  setiger  5,  becoming  rather  low  and  wide  on  more  posterior 
segments.  With  capillary  setae  in  both  notopodia  and  neuropodia; 
beginning  on  setiger  7,  neuropodial  capillary  setae  mostly  replaced 
by  hooded  bidentate  crotchets.  In  more  posterior  setigers  (last  20 
or  so),  notopodia  conical,  lacking  branchiae  or  lamellae,  with  conical 
bundle  of  awl-shaped  setae  scarcely  surpassing  notopodial  tip  and 
small  bundle  of  capillary  setae;  neuropodia  with  two  or  three  crotchets 
and  few  capUlary  setae.  Anal  disc  with  complete  middorsal  notch, 
shorter  midventral  and  midlateral  notches,  thus  4-lobed.  Color:  In 
alcohol:  Without  color  or  grayish. 

Remarks. — The  type  of  P.  brachycephala  Hartman  from  central 
California  was  examined  and  is  herein  referred  to  P.  caulleryi. 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  up  to  8 
miles  from  shore,  16.7-75.5  fms.,  on  bottoms  of  stones  and  various 
combinations  of  mud,  gravel,  stones,  rocks,  large  perforated  rocks, 
with  bryozoans  and  hydroids  (8  stations,  16  specimens).  West  Coast 
North  America:  Washington  and  Puget  Sounds,  low  water  to  50 
fms.,  Pettibone. 

Distribution. — Arctic  Alaska  to  central  California;  Ireland,  Great 
Britain  to  France.     In  low  water  to  75.5  fathoms. 

Polydora  larva 

A  larva  of  a  Polydora  was  taken  in  plankton  August  2,  1948.  This 
was  perhaps  the  larva  of  P.  caulleryi  Mesnil  since  it  was  the  only 
species  of  Polydora  found  in  the  Point  Barrow  region.  Prostomium 
rounded,  with  two  palps,  and  with  black  chroma tophores.  Fifth 
setigerous  segment  modified,  with  short,  thick,  slightly  curved  acicular 
setae;  rest  of  21  setigerous  segments  with  long  larval  capillary  setae. 
Pygidium  a  rounded  disc,  notched  middorsaUy,  with  black  chromato- 
phores.  Black  chroma  tophores  arranged  transversely  on  setigers  2-5, 
then  in  two  longitudinal  lines  on  each  side  of  middorsal  line,  extending 
to  anal  cup. 


282  PROCEEDINGS    OF   THE   NATIONAL  MUSEUM  tol.io3 

C^nus  Prionospio  Malmgren,  1867 
Prionospio  malmgreni  Clapar^de,  1868 

Figure  32,  i-k 

Prionospio  malmgreni  ClaparMe,  1868,  p.  333,  pi.  22,  fig.  3  (fide  Fauvel,  1927). — 
Ehlers,  1913,  p.  511.— Fauvel,  1927,  p.  61,  fig.  21,  a-e.— Annenkova,  1937, 
p.  171;  1938,  p.  174.— Hartman,  1948,  p.  36.— Wesenberg-Lund,  1951, 
p.  70. — Berkeley  and  Berkeley,  1952,  p.  29,  figs.  54,  55. 

Spiophanes  tenuis  Verrill,  1879b,  p.  176. 

Prionospio  tenuis  Verrill,  1882,  p.  370. — Hartman,  1944a,  pp.  336,  340,  ?  pi.  19, 
fig.  7;  not  1945,  p.  32  (  =  P.  treadwelli  Hartman,  1951). 

Description. — Length  up  to  25  mm.,  width  1  mm.  Prostomium 
dilated  anteriorly,  with  anterior  border  rounded,  narrowing  posteriorly, 
extending  in  obtuse  crest  to  setiger  2  (to  4) .  Four  eyes,  two  smaller, 
two  much  larger  (may  be  lacking) .  Two  long  palps,  usually  missing 
(in  alcohol).  A  short,  eversible,  somewhat  lobulated  proboscidial 
region.  Branchiae  four  pairs,  on  setigers  2-5,  completely  separated 
from  dorsal  lamellae,  pairs  1  and  4  longer,  pinnate,  somewhat  decid- 
uous, pairs  2  and  3  shorter,  smooth,  triangular.  Dorsal  postsetal 
lamellae  of  first  setiger  small,  much  larger  and  triangular  on  next 
four  setigers,  then  diminishing  in  size,  becoming  reduced  beginning 
on  setigers  8  or  9;  ventral  postsetal  lamellae  oval,  rounded.  Hooded, 
multidentate  crotchets  beginning  about  setigers  12-16  in  neuropodia 
and  about  setiger  40  in  notopodia.  Beginning  on  setiger  7,  slightly 
elevated  transverse  folds  between  dorsal  lamellae,  continuing  on  next 
6-14  segments.  Pygidium  with  longer,  unpaired  dorsal  cirrus  and 
two  smaller  lateral  ones.     Without  color  in  alcohol. 

Remarks. — The  types  of  Prionospio  tenuis  Verrill  from  Cape  Cod 
Bay  were  examined  and  are  herein  referred  to  P.  malmgreni.  There 
are  four  pairs  of  branchiae,  the  first  and  fourth  pairs  pinnate,  the  sec- 
ond and  third  pairs  shorter,  smooth. 

New  records, — Arctic  Alaska:  Off  Point  Barrow  base,  4  miles  from 
shore,  29  fms.,  on  bottom  of  gravel,  small  stones  (1  station,  2  speci- 


FiGURE  32. — Orbiniidae:  a,  Scoloplos  armiger,  dorsal  view  anterior  end;  i,  same,  parapodium 
from  thoracic  region,  posterior  view;  c,  same,  parapodium  from  abdominal  region,  anterior 
view;  d,  ty  same,  crenulate  neurosetae  from  thoracic  region.  Spionidae:/,  Spio  filicornis, 
dorsal  view  anterior  end  (palps  missing);  g,  same,  parapodium  from  anterior  region, 
anterior  view;  h,  same,  hooded  bidentate  crotchet;  i,  Prionospio  malmgreni,  dorsal  view 
anterior  end;  /,  same,  branchiae  on  setigers  2-S  (one  of  each  pair  shown);  k,  same,  hooded 
multidentate  crotchet;  /,  Polydora  caulleryi,  dorsal  view  anterior  end  (palps  missing); 
m,  same,  parapodium  of  modified  fifth  setigerous  segment;  «,  same,  special  hooks  from 
setiger  5;  o,  same,  notopodium  of  posterior  setiger,  with  conical  bundle  awl-shaped 
setae;  p,  same,  anal  disc  of  pygidium,  lateral  view;  q,  same,  hooded  bidentate  crotchet, 
f,  Ntrinides  sp.,  dorsal  view  anterior  end;  s,  same,  parapodium  from  anterior  region. 
(For  explanation  of  symbols,  see  p.  210.) 


MARINE    POLYCHAETE    WORMS — PETTIBONE  283 


P  Q 

Figure  32. — For  explanation  see  facing  page. 


261112—54 6 


284  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  103 

mens).  West  Coast  North  America:  Puget  Sound,  Washington, 
low  water,  mud,  Pettibone.  East  Coast  North  America:  Off 
Labrador,  40  fms,,  Blue  Dolphin  Expedition,  1949;  off  New  Hamp- 
shire and  Massachusetts,  25-48  fms.,  U.  S.  Fish  Commission. 

Distribution. — Arctic  Alaska  to  Washington;  north  Japan  Sea; 
Iceland,  Norway  to  Mediterranean;  Labrador  to  Massachusetts; 
South  Africa.     In  low  water  to  250  fathoms;  surface. 

Genus  Spio  Fabricius,  1785 
Spiofilicornis  (Miiller,  1776) 

Figure  32,  f-h 

Nereis  filicornis  Miiller,  1776,  p.  218. 

Spio  filicornis  Malmgren,  1867,  p.  91,  pi.  1,  fig.  1. — Eliason,  1920,  p.  40,  figs. 
7-9.— Fauvel,  1927,  p.  43,  fig.  15,  a-g.— Augener,  1928,  p.  738,  pi.  11,  fig.  7.— 
Annenkova,  1937,  p.  169;  1938,  p.  173.— Hartman,  1941a,  p.  293;  1948,  p. 
36.— Berkeley  and  Berkeley,  1943,  p.  130.— Thorson,  1946,  p.  93,  fig.  46.— 
Zatsepin,  1948,  p.  132,  pi.  32,  fig.  11.— Wesenberg-Lund,  1950a,  p.  30; 
1950b,  p.  76;  1951,  p.  71. 

Spio  limicola  Verrill,  1879,  p.  176. 

Spio  filicornis  var.  pacifica  Berkeley  and  Berkeley,  1936,  p.  475;  1952,  p.  25,  figs. 
47,  48. 

Description. — Up  to  30  mm.  long,  2  mm.  wide.  Prostomium  bell- 
shaped  to  conical,  widest  anteriorly,  rounded,  tapered  posteriorly  to 
short  crest  extending  on  first  two  setigers.  Four  eyes  in  trapezoidal 
arrangement,  anterior  pair  larger.  Two  palps  missing  (in  alcohol). 
A  short,  eversible,  somewhat  lobulated  proboscidial  region.  Branchiae 
begin  on  setiger  1,  continuing  to  posterior  end,  simple,  straplike. 
Dorsal  postsetal  lamellae  of  more  anterior  segments  short,  wide,  with 
branchiae  fused  to  upper  sides;  in  more  posterior  segments,  lamellae 
cirriform,  separate  from  branchiae.  Ventral  postsetal  lamellae 
rounded,  not  notched.  Capillary  setae  only  in  notopodia.  Neuropodia 
with  hooded,  bidentate  crotchets  beginning  on  setigers  9-18,  about  six 
in  number  (5-10).  Pygidium  with  four  cirri?  Color:  In  alcohol: 
Yellowish  white  with  dark  brown  pigment  on  peristomial  region  and 
along  each  side  of  anterior  fourth  of  body. 

Remarks. — The  types  of  Spio  limicola  Verrill  from  Cape  Cod  Bay 
were  examined  and  are  herein  referred  to  S.  jilicornis.  S.  limicola 
differs  from  S.  setosa  Verrill,  to  which  it  has  been  referred  (Hart- 
man,  1942,  p.  63),  in  the  number  of  neuropodial  hooded  crotchets, 
6-9  in  S.  limicola  and  about  16  in  ^S.  setosa. 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  up  to  8 
miles  from  shore,  36-75.5  fms.,  on  various  combinations  of  gravel, 
rocks,  stones,  large  perforated  rocks  (2  stations,  2  specimens) .  West 
Coast  North  America:  Washington  Sound,  low  water,  Pettibone. 
East  Coast  North  America:  Off  Massachusetts,  15-31  fms.,  U.  S. 
Fish  Commission. 


MARINE    POLYCHAETE    WORMS — PETTIBONE  285 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian  and 
Alaskan  Arctic,  Davis  Strait,  Greenland,  Spitsbergen,  Novaya  Zemlya. 
Also  Iceland,  Faroes,  Norway  to  France;  Hudson  Bay  to  Massachu- 
setts; Bering  Sea  to  central  California;  Japan.  In  low  water  to  211 
fathoms;  sm-face. 

Genus  Nerinides  Mesnil,  1896 
Nerinides  sp. 

Figure  32,  r,  s 

Two  small  specimens  obtained  from  plankton  10  feet  from  shore, 
agreeing  in  some  respects  with  the  description  of  N.  cantabra  Rioja 
(Fauvel,  1927,  p.  31).  Length  16  mm.,  width  1  mm.,  segments  57. 
Prostomium  rounded  anteriorly,  tapering  posteriorly,  with  middorsal 
crest  extending  on  first  setiger,  with  four  eyes,  one  pair  larger,  a 
raised  area  just  posterior  to  eyes  (median  antenna?),  with  two  large, 
thick  palps  (fig.  32,  r).  Dorsal  and  ventral  postsetal  lamellae  distinct, 
dorsal  ones  may  be  slightly  undulate,  ventral  ones  entire,  not  notched 
(fig.  32,  s).  Branchiae  begin  on  setiger  2,  continuing  to  near  posterior 
end,  fused  to  upper  part  of  dorsal  lamellae.  Body  with  low  transverse 
dorsal  ridges  between  bases  of  branchiae.  First  setiger  with  both 
dorsal  and  ventral  setae.  Notopodia  with  only  capillary  setae,  being 
extra  long  in  posterior  region.  Neuropodia  with  capillary  setae  and 
hooded  bidentate  crotchets  beginning  around  setiger  20.  Pygidium 
with  4-lobed  membranous  appendage.    Without  color  in  alcohol. 

Family  Cirratulidae 

Body  more  or  less  cylindrical,  attenuated  at  both  ends,  with 
numerous,  short,  compact,  nearly  similar  segments.  Prostomium 
small,  usually  without  appendages,  with  or  without  eyes.  Pair  of  large 
grooved  spioniform  palps  (fig.  33,  d)  or  with  more  or  less  numerous 
grooved  tentacular  fiilaments  inserted  on  dorsal  surface  of  an  anterior 
segment  (fig.  33,  a).  Branchiae  simple,  filamentous,  long,  con- 
tractile, inserted  above  notopodia  on  few  to  many  segments  (may 
be  absent).  Parapodia  biramous,  lobes  scarcely  projecting  (fig.  33, 
6).  Setae  usually  simple,  some  capillary,  others  acicular  crotchets 
(fig.  33,  c) .  Without  dorsal,  ventral,  or  anal  cirri.  Proboscis  smooth, 
unarmed.  May  have  epitokous  pelagic  form  at  time  of  reproduction. 
Usually  in  mud;  some  (Dodecaceria)  in  calcareous  matrix. 

Represented  by  two  genera  and  two  species  from  Point  Barrow. 
Both  genera  have  prostomium  conical,  buccal  segment  and  two  fol- 
lowing segments  achaetous,  more  or  less  fused  to  prostomium,  with 
lateral  filamentous  branchiae  beginning  just  anterior  to  first  setigerous 
segment,  continuing  on  large  number  of  segments. 


286  PROCEEDINGS    OF   THE    NATIONAL  MUSEUM  vol.108 

Key  to  the  genera  of  Cirratulidae  from  Point  Barrow 

1.  With  few  to  numerous,  grooved  tentacular  filaments  just  anterior  to  first 
setigerous  segment,  subequal  to  branchiae  (without  2  palps;  fig.  33,  a). 

Cirratulus   (p.   286) 

With  2  large  grooved,  spioniform  palps  just  anterior  to  first  setigerous  segment 

(without  tentacular  filaments;  palps  deciduous;  may  be  missing  in  epitoke; 

fig.  33,  d) Chaetozone  (p.  287) 

Genus  Cirratulus  Laxuark,  1801 
Cirratulus  cirratus  (Miiller,  1776) 

Figure  33,  a-c 

Lumbricus  cirratus  Miiller,  1776,  p.  215. 

Cirratulus  cirratus  Malmgren,  1867,  p.  95. — Chamberlin,  1920,  p.  20. — Eliason, 
1920,  p.  57.— Fauvel,  1927,  p.  94,  fig.  33,  a-g;  1933,  p.  46;  1936,  p.  72.— 
Augener,  1928,  p.  750,  pi.  11,  fig.  9.— Monro,  1930,  p.  154;  1936,  p.  161.— 
Annenkova,  1931,  p.  203;  1932,  p.  136;  1934,  p.  322;  1937,  p.  173;  1938,  p. 
181.— Berkeley  and  Berkeley,  1942,  p.  197;  1943,  p.  130;  1952,  p.  31,  figs. 
58,  59.— Hartman,  1944a,  pp.  334,  341;  1944b,  p.  263;  1948,  p.  37.— Gorbunov, 
1946,  p.  39.— Zatsepin,  1948,  p.  133,  pi.  32,  fig.  18.— Wesenberg-Lund, 
1950a,  p.  33;  1950b,  p.  81;  1951,  p.  74.— Hartman  and  Reish,  1950,  p.  34. 

Cirratulus  robustus  Johnson,  1901,  p.  423,  pi.  14,  figs.  149,  150. — Rioja,  1941, 
p.  728.— Berkeley  and  Berkeley,  1942,  p.  197. 

Cirratulus  cingulatus  Johnson,  1901,  p.  422,  pi.  14,  figs.  145-148. — Rioja,  1941, 
p.  729. 

Cirratulus  spectabilis  Berkeley  and  Berkeley,  1952,  p.  32. 

Description. — Length  up  to  300  ^mm.,  width  5  mm.  Prostomium 
variable  in  shape:  Short,  broad,  bluntly  conical;  almost  square- 
shaped;  or  tapered  gradually,  then  abruptly.  Prostomium  with 
usually  four  or  five  (2-8)  eyes  on  each  side  in  obliquely  transverse 
row  or  may  be  united  in  almost  solid  arc.  Buccal  segment  more  or 
less  fused  with  prostomium,  two  following  achaetous  segments  more 
or  less  distinct.  Tentacular  filaments  anterior  to  first  setigerous 
segment,  in  two  groups  of  2-24  in  obliquely  transverse  rows,  long, 
filamentous,  grooved  longitudinally.  Branchiae  begin  just  anterior  to 
first  setigerous  segment,  lateral  to  group  of  tentacular  filaments, 
continuing  through  greater  part  of  body,  anteriorly  inserted  close  to 
upper  part  of  notopodia,  then  gradually  shifting  more  dorsally  and 
then  sUghtly  more  ventrally.  Parapodia  consisting  of  slightly  pro- 
jecting tori  common  to  both  no  to-  and  neuropodia,  with  capillary 
setae  in  both  lobes,  with  additional  acicular  crotchets  first  appearing 
in  neuropodia  and  more  posteriorly  in  notopodia  also.  Anus  sub- 
dorsal. Color:  In  life:  Orange  yellow  with  red  branchiae;  dark 
orange  brown  with  orange-red  branchiae.  In  alcohol:  Yellowish 
brown  or  without  color. 

New  records. — Arctic  Alaska:  Eluitkak  Pass,  Elson  Lagoon  near 
Point  Barrow,  5-6.6  fms.;  off  Point  Barrow  base,  up  to  8  miles  from 


MARINE    POLYCHAETE    WORMS — PETTIBONE  287 

shore,  13.3-75.5  fms.,  on  bottoms  of  stones,  and  various  combinations 
of  mud,  pebbles,  rock,  stones,  gravel,  large  perforated  rocks,  with 
bryozoans,  embedded  in  mud  at  bases  of  barnacles  and  bryozoans  as 
Eucratea  loricata  (19  stations,  183  specimens).  Siberia:  Plover  Bay, 
10-35  fms.,  Dall,  1880.  Bering  Sea:  Albatross  station,  Nikolski, 
Bering  Island,  1892;  St.  George  Island,  Pribilofs,  2  miles  off  shore, 
40  fms.,  Hanna,  1913;  Atka  Island,  Aleutians,  L.  M.  Turner,  1879. 
West  Coast  North  America:  Strait  of  Juan  de  Fuca,  Washington 
and  Puget  Sounds,  low  water,  Pettibone.  East  Coast  North  Amer- 
ica: Off  Labrador,  95  fms..  Blue  Dolphin  Expedition,  1949;  oil  Maine, 
Massachusetts,  low  water  to  67  fms.,  U.  S.  Fish  Commission. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian,  Alaskan, 
and  Canadian  Arctic,  Greenland,  Spitsbergen,  Barents  Sea,  Novaya 
Zemlya.  Also  Iceland,  Faroes,  Norway  to  France,  Canary  Islands; 
Hudson  Bay  to  Massachusetts;  Bering  Sea  to  Mexico;  north  Japan 
Sea  to  Japan,  Manchuria.  Southern  latitudes:  Falkland  Islands, 
Magellan  Straits,  South  Georgia,  Kerguelen.  In  low  water  to  1,611 
fathoms. 

Genus  Chaetozone  Malmgren,  1867 

Chaetozone  setosa  Malmgren,  1867 

Figure  33,  d 

Chaetozone  setosa  Malmgren,  1867,  p.  96,  pi.  14,  fig.  84. — Th^el,  1879,  p.  54,  pi. 
4,  figs.  49-51.— Fauvel,  1914,  p.  217,  pi.  20,  fig.  1;  1927,  p.  101,  fig.  35; 
1934a,  p.  47.— Eliason,  1920,  p.  57. — Augener,  1928,  p.  750. — Gustafson, 
1936,  p.  8.— Monro,  1937,  p.  301.— Annenkova,  1937,  p.  174;  1938,  p.  182.— 
Berkeley  and  Berkeley,  1942,  p.  197;  1952,  p.  35,  fig.  63.— Hartman,  1944a, 
pp.  334,  341.— Zatsepin,  1948,  p.  133,  pi.  32,  fig.  16.— Wesenberg-Lund, 
1950a,  p.  34;  1950b,  p.  81;  1951,  p.  75. 

Description. — Length  up  to  25  mm.,  width  2  mm.  Prostomium 
conical,  acutely  pointed,  without  eyes.  Buccal  segment  and  two 
following  achaetous  segments  more  or  less  fused;  in  specimens  from 
Point  Barrow,  buccal  segment  fused  with  prostomium,  second  and 
third  segments  fused  dorsally.  Paired  palps  large,  long,  grooved, 
inserted  just  anterior  to  first  setigerous  segment,  deciduous,  often 
missing  on  preserved  specimens  and  missing  when  in  epitokous  phase. 
Branchial  filaments  begin  just  anterior  to  first  setigerous  segment, 
immediately  posterior  to  palps,  continuing  on  large  number  of  anterior 
segments,  lacking  on  posterior  region,  readily  deciduous,  inserted  just 
posterior  to  upper  part  of  parapodial  tori.  Parapodia  are  slightly 
projecting  tori  common  to  both  noto-  and  neuropodia.  In  anterior 
region,  both  lobes  with  long,  delicate  capillary  setae;  more  posteriorly, 
notopodia  with  still  longer  capillary  setae,  neuropodia  with  shorter, 
stouter  capillary  setae  and  short,  unidentate  acicular  crotchets;  more 
posteriorly,  with  acicular  crotchets  in  both  noto-  and  neuropodia. 


288  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  TOL.108 

In  epitokous  phase,  dorsal  setae  in  anterior  region  much  longer;  ia 
posterior  region,  parapodial  tori  more  projecting,  dorsal  and  ventral 
acicular  crotchets  forming  semicircular,  fan-shaped  group,  almost  en- 
circHng  body.  Pygidium  with  small,  rounded  to  conical  lobe,  ventral 
and  posterior  to  anus  (like  a  cover  for  anal  opening).  Color:  In 
alcohol:  Light  to  dark  gray,  brownish,  or  iridescent  bluish  black. 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  up  to  12.1 
miles  from  shore,  20-123.5  fms.,  on  various  combinations  of  mud, 
stones,  gravel,  rocks,  large  perforated  rocks,  mass  of  worm  tubes,  on 
surface  of  tunicates  (12  stations,  29  specimens).  East  Coast  North 
America:  Off  Labrador,  95-125  fms..  Blue  Dolphin  Expedition,  1949; 
off  Maine,  New  Hampshire,  Massachusetts,  11-58  fms.,  U.  S.  Fish 
Commission. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian  and 
Alaskan  Arctic,  Davis  Strait,  Greenland,  Spitsbergen,  Franz  Josef 
Land,  Barents  Sea,  Novaya  Zemlya.  Also  Iceland,  Faroes,  Norway 
to  Canary  Islands,  Mediterranean,  Adriatic,  Gulf  of  Aden;  Labrador 
to  Massachusetts;  Bering  Sea  to  British  Columbia;  north  Japan  Sea. 
Southern  latitudes:  Falkland  Islands,  Magellan  Straits,  Kerguelen. 
In  low  water  to  1,333  fathoms;  surface. 

Family  Flabelligeridae  (Chloraemidae) 

Body  subcylindrical  or  subfusiform,  with  short  similar  segments 
covered  with  papillae,  often  with  adherent  sand  grains,  may  be 
encased  in  thick  mucous  mantle.  Prostomium  and  buccal  segment 
fused,  with  two  thick  grooved  palps,  two  groups  of  filiform  cephalic 
branchiae,  and  more  or  less  developed  buccal  tube  around  mouth, 
all  of  which  may  be  partially  or  completely  retracted  within  anterior 
setigers,  with  or  without  setae  of  anterior  setigers  directed  anteriorly 
forming  a  cephaUc  cage.  Parapodia  biramous;  notosetae  simple, 
capillary,  ringed;  neurosetae  simple  or  pseudocompound,  capillary 
or  hooked.  Without  dorsal,  ventral,  or  anal  cirri.  Anus  terminal. 
Usually  in  muddy  sand. 

Represented  by  two  genera  and  three  species  from  Point  Barrow. 

Key  to  the  genera  of  Flabelligeridae  from  Point  Barrow 

1.  Body  covered  with  long  pedunculate  papillae  (fig.  33,  g),  usually  encased  in 
thick  mucous  mantle.  Fused  prostomium  and  buccal  segment  partially 
retractile  within  first  setigers  and  more  or  less  hidden  by  cephalic  cage  formed 
by  setae  of  first  setiger,  the  setae  long,  directed  anteriorly,  almost  completely 

encircling  segment Flabelligera  (p.  289) 

Body  covered  with  short  papillae,  usually  encrusted  with  sand,  not  encased 
in  mucous  mantle.  Fused  prostomium  and  buccal  segment  completely 
retractile  within  anterior  setigers,  may  be  completely  hidden;  without 
cephalic  cage  formed  by  anterior  setigers;  when  buccal  segment,  etc.  are 
retracted,  anterior  end  is  trilobed  (fig.  33,  h) Brada  (p.  290) 


MARINE    POLYCHAETE    WORMS — PETTIBONE  289 

Genus  Flabelligera  Sars,  1829 

Flabelligera  affinis  Sars,  1829 
Figure  33,  e~g 

Flabelligera  affinis  Sars,  1829,  p.  31,  pi.  3,  fig.  16  (fide  Malmgren,  1867). — Malmgren, 
1867,  p.  83.— Chamberlin,  1920,  p.  21.— Eliason,  1920,  p.  62.— Fauvel,  1927, 
p.  113,  fig.  40,  a-f.— Augener,  1928,  p.  768.— Monro,  1930,  p.  160,  fig.  63; 
1936,  p.  164.— Annenkova,  1937,  p.  174;  1938,  p.  184.— Berkeley  and  Berkeley, 
1943,  p.  130.— Hartman,  1944a,  pp.  335,  341,  pi.  33,  figs.  12,  13  (as  ?Brada 
setosa  Verrill). — Gorbunov,  1946,  p.  39. — Zatsepin,  1948,  p.  134,  pi.  33,  fig. 
1.— St0p-Bowitz,  1948a,  p.  8,  fig.  1;  1948b,  p.  30.— Wesenberg-Lund,  1950a, 
p.  35;  1950b,  p.  82;  1951,  p.  76. 

Flabelligera  infundibularis  Johnson,  1901,  p.  417,  pi.  12,  figs.  124-127. — Hartman, 
1938c,  p.  14;  1948,  p.  40.— Berkeley  and  Berkeley,  1952,  p.  7,  figs.  1-4. 

Flabelligera  infundibularum  Moore,  1923,  p.  223. 

Flabelligera  infundibuliformis  Hartman  and  Reish,  1950,  p.  35. 

Description. — Length  up  to  130  mm.,  width  including  mantle  12-15 
mm.,  segments  up  to  7 1 .  Body  rounded,  attenuated  shghtly  anteriorly 
and  more  so  posteriorly.  Body  covered  with  numerous  papillae 
which  are  elongate,  thin,  flexible,  usually  somewhat  coiled,  with  larger 
bulbous  tips  quite  variable  in  shape.  Body  usually  covered  with 
thick,  mucous,  transparent  mantle  (may  be  missing  in  smaU  speci- 
mens or  specimens  appearing  in  surface  waters)  which  is  penetrated 
by  the  pedunculate  papillae.  Fused  prostomium  and  buccal  segment 
retractile  within  anterior  setiger,  with  pair  of  stout,  grooved  palps, 
two  groups  of  numerous  filiform  branchiae,  two  pairs  of  eyes  (almost 
coalesced),  and  somewhat  protruding  buccal  siphon  around  ventral 
mouth  opening.  First  setigerous  segment  with  wide,  funneUike 
depression  into  which  buccal  siphon,  etc.,  can  mostly  be  withdrawn; 
with  circlet  of  elongated,  ringed  setae  around  margin  and  directed 
forward  or  spread  fimnellike,  forming  cephalic  cage;  with  numerous 
elongate  papillae  having  slender  straight  peduncles  and  fusiform  tips. 
Notopodia  conical,  diagonally  truncate  lobes,  with  capillary  ringed 
setae,  with  elongate  papillae  on  posterior  side  that  may  be  up  to  three- 
fourths  length  of  notosetae  (similar  in  shape  to  those  around  cephalic 
cage;  different  from  those  on  body).  Neuropodia  conical,  each  with 
usually  one  (sometimes  two)  large,  hooked,  amber-colored  pseudo- 
compoimd  seta,  ringed  basaUy,  hooked  tips  dark  to  black.  When 
mucous  mantle  is  present,  setae  almost  completely  embedded  and  the 
hooked  tips  of  neurosetae  and  capillary  tips  of  notosetae  only  ex- 
tending beyond  mantle.  Pygidium  a  fleshy,  rounded  rim;  anus 
subdorsal.  Color:  In  alcohol:  Without  color  or  slightly  reddish 
brown. 

New  records. — Arctic  Alaska:  West  side  Elson  Lagoon  near  en- 
trance to  small  lagoon  to  west,  1.2  fms.;  Point  Barrow  base,  washed 
ashore;  off  Point  Barrow  base,  18.3-123.5  fms.,  on  various  combina- 


290  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  los 

tions  of  mud,  stones,  gravel,  rocks,  with  mass  of  worm  tubes  (12 
stations,  15  specimens).  Bering  Sea:  Albatross  Sta.  3274,  55°34' 
N.,  162°31'  W.,  19  fms.,  and  station  at  Unalaska,  Aleutians,  1890; 
IluUiuk  Harbor,  Unalaska,  Dall,  1871 .  West  Coast  North  America  : 
Washington  and  Puget  Sounds,  10  fms.  and  surface,  Pettibone. 
Canadian  Arctic:  Dobbin  Bay,  east  Ellesmere  Land,  79°36'  N., 
73°35'  W.,  16  fms.,  W.  H.  Littlewood,  1950.  Foxe  Basin,  Baffin 
Island,  66°83'  N.,  80°07'  W.,  Bartlett,  1927.  West  Greenland: 
Between  Capes  Alexander  and  Chalon,  25-40  fms.,  Bartlett,  1937. 
East  Coast  North  America:  Off  Labrador,  10-30  fms.,  Blue 
Dolphin  Expeditions,  1949,  1950;  Grand  Manan,  Maine,  New 
Hampshire,  Massachusetts,  3-90  fms.,  U.  S.  Fish  Commission. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian,  Alaskan, 
and  Canadian  Arctic,  Greenland,  Spitsbergen,  Franz  Josef  Land, 
Barents  Sea,  White  Sea,  Novaya  Zemlya,  Kara  Sea.  Also  Iceland, 
Faroes,  Norway  to  France,  Mediterranean;  Hudson  Bay  to  Mas- 
sachusetts; Bering  Sea  to  southern  California;  north  Japan  Sea. 
Southern  latitudes:  Falkland  Islands,  Magellan  Straits,  South 
Georgia,  South  Africa,  Australia.  In  low  water  to  889  fathoms; 
surface. 

Genus  Brada  Stimpson,  1854 

Key  to  the  species  of  Brada  from  Point  Barrow 

1.  Body  covered  with  filiform  papillae,  often  encrusted  with  sand  grains,  with 
tips  of  papillae  bare.     Notopodia  distinct,  with  2-5  capillary  setae  per  lobe. 

B.  villosa 
Body  covered  with  low  globular  papillae,  often  encrusted  with  a  thin  layer  of 
sand  grains.    Notopodia  indistinct,  with  0-2  very  delicate  capillary  setae. 

B.  inhabilis 

Brada  villosa  (Rathke,  1843) 

Siphonostoma  villosum  Rathke,  1843,  p.  215,  pi.  11,  figs.  11,  12. 

Brada  villosa  Malmgren,  1867,  p.  84.— Eliason,  1920,  p.  62.— Chamberlin,  1920, 

p.  22.— Fauvel,  1927,  p.  121,  fig.  43,  e-1;  1934a,  p.  49.— Monro,  1930,  p.  161; 

1937,  p.   303.— Annenkova,    1937,   p.    175;   1938,   p.    185.— Okuda,   1937b, 

p.  53,  pi.  2,  fig.  D.— Berkeley  and  Berkeley,  1943,  p.  130;  1952,  p.  7,  fig.  5.— 

Zatsepin,   1948,  p.   136,  pi.  33,  fig.  4.— St0p-Bowitz,   1948a,  p.  33,  fig.  9; 

1948b,  p.  39,  fig.  15. — Hartman  and  Reish,  1950,  p.  35. — Wesenberg-Lund, 

1950a,  p.  35;  1950b,  p.  85;  1951,  p.  78,  fig.  7. 
Brada  setosa  Verrill,  1873,  pp.  431,  434,  508,  606;  1881,  p.  302,  pi.  9,  fig.  4.— 

Sumner,  Osburn  and  Cole,  1913,  p.  630. 
Trophonia  rugosa  Hansen,  1882,  p.  38,  pi.  7,  figs.  9-12. 
Trophonia  arctica  Hansen,  1882,  p.  39,  pi.  7,  figs.  17-20. 
Brada  granulata  Murdoch,  1885,  p.  155  (not  B.  granulata  Malmgren,  1867). 
Brada  pilosa  Moore,  1906b,  p.  231,  pi.  10,  figs.  14-17. 
Stylaroides  pluribranchiata  Moore,  1923,  p.  222. 
Brada  rugosa  St0p-Bowitz,  1948a,  p.  37,  fig.  10;  1948b,  p.  41. 


MARINE    POLYCHAETE    WORMS — PETTIBONE  291 

Description. — Length  28-60  mm.,  width  5-9  mm.,  setigers  23-35. 
Body  fusiform,  slightly  flattened  ventrally,  convex  dorsally,  attenu- 
ated and  more  or  less  elongated  posteriorly,  having  a  characteristic 
rough,  hirsute  aspect,  covered  with  cylindrical,  fusiform  to  claviform 
papillae,  longer  dorsally  than  ventrally,  usually  encrusted  with  sand 
grains  more  basally,  with  distal  tips  bare;  there  are  all  gradations: 
Without  encrusting  sand  (looks  "hairy"),  with  scattered  sand  grains, 
and  with  rather  thick  layer  of  sand  grains,  usually  fewer  on  antero- 
ventral  part.  Underlying  surface  nodular  and,  in  larger  specimens, 
outer  papillate  layer  may  be  worn  off,  the  surface  appearing  rugose, 
with  low  mounds.  Retractile  prostomium  and  buccal  region  with  two 
groups  of  numerous  filiform  branchiae  borne  on  pair  of  dorsolateral 
bosses,  pair  of  thick  grooved  palps,  and  ventral  eversible  buccal  siphon 
around  mouth.  Setae  of  first  setigerous  segment  8-10  per  bundle, 
slightly  more  elongate  than  following,  directed  anteriorly.  Notopodia 
and  neuropodia  distinct,  with  long  fusiform  papillae  grouped  around 
setae.  Notosetae  long,  capillary,  ringed,  2-5  in  number.  Neurosetae 
beginning  on  setiger  2,  large,  3-6  in  number,  amber-colored,  ending  in 
sharp,  transparent,  and  fragile  tips.  Pair  of  nephridial  papillae  near 
anterior  border  of  ventral  side  of  setiger  5.  Color:  In  life  and  in 
alcohol:  Brownish,  dull  sandy-mud. 

Remarks. — The  types  of  Brada  setosa  Verrill  from  Massachusetts, 
B.  pilosa  Moore  from  Alaska,  and  Stylaroides  plurihranchiata  Moore 
from  California  were  examined  and  are  hereui  referred  to  B.  villosa. 
The  excessive  "hairiness"  of  B.  pilosa  appears  to  be  a  variable  charac- 
ter; there  may  be  varying  numbers  of  papillae  as  well  as  all  gradations 
in  the  amount  of  encrusting  sand  grains.  Two  large  specimens  washed 
ashore  at  Point  Barrow  were  at  first  identified  as  B.  rugosa,  following 
St0p-Bowitz  (1948);  they  agree  in  the  large  size  (23-24  setigers,  28-30 
mm.  long,  5-7  mm.  wide),  the  almost  complete  absence  of  papulation, 
and  the  thick  rugose  surface;  the  outer  surface,  however,  appears  to  be 
worn  off.  There  are  some  remains  of  the  papillated  surface  on  the 
parapodial  lobes.  Thus  B.  rugosa  (including  T.  arctica)  is  herein 
referred  to  B.  villosa,  where  it  has  been  referred  earlier  by  Fauvel  and 
others. 

New  records. — Arctic  Alaska:  Point  Barrow  base,  washed  ashore 
September  22,  1949  (22  specimens),  August  10,  1950  (2  specimens);  off 
Point  Barrow  base,  up  to  4  mUes  from  shore,  27-29  fms.,  on  various 
combinations  of  mud,  gravel,  stones,  rocks  (2  stations,  2  specimens), 
Bering  Sea:  62°15'  N.,  167°48'  W.,  Stoney,  1884;  Albatross  Sta.  3337, 
53°55'  N.,  163°26'  W.,  280  fms.,  1890.  West  Coast  North  America: 
Washington  Sound,  Pettibone;  Albatross  Sta.,  northwest  West  Point, 
Elliot  Bay,  Seattle,  1914.    East  Coast  North  America:  Off  Nova 


292  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  voi-.ios 

Scotia,  Maine,  Massachusetts,  Rhode  Island,  6-499  fms.,  U.  S.  Fish 
Commission. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian,  Alaskan, 
and  Canadian  Arctic,  Greenland,  Spitsbergen,  Barents  Sea,  Novaya 
Zemlya,  Kara  Sea.  Also  Iceland,  Faroes,  Norway  to  Spain,  Mediter- 
ranean, Adriatic,  south  Arabian  Coast;  Hudson  Bay  to  Rhode  Island; 
Bering  Sea  to  southern  California;  north  Japan  Sea  to  Japan;  South 
Atlantic:  South  Orkney  and  South  Shetland  Islands.  In  low  water 
to  853  fathoms. 

Brada  inhabilis  (Rathke,  1843) 

FiGUEE  33,  h 

Siphonostoma  inhahile  Rathke,  1843,  p.  218,  pi.  11,  fig.  13. 

Brada  sublaevis  Stimpson,  1854,  p.  32.— Verrill,  1881,  pp.  289,  304.— Webster 
and  Benedict,  1887,  p.  731. 

Brada  granulata  Malmgren,  1867,  p.  85,  pi.  12,  fig.  71.— Th^el,  1879,  p.  52.— 
not  Murdoch,  1885,  p.  155  (  =  S.  villosa). — Annenkova,  1937,  p.  176;  1938, 
p.  186.— Hartman,  1944a,  pp.  334,  341;  1948,  p.  41.— Zatsepin,  1948,  p. 
135,  pi.  33,  fig.  3.— Wesenberg-Lund,  1950a,  p.  36;  1950b,  p.  85. 

Brada  granosa  Treadwell,  1937,  p.  32  (not  B.  granosa  Stimpson,  1854). 

Brada  inhabilis  St0p-Bowitz,  1948a,  p.  40,  fig.  11;  1948b,  p.  42,  fig.  16.— Wesen- 
berg-Lund, 1951,  p.  80. 

Description. — Length  30-60  mm.,  width  8-12  mm.,  segments  23-26. 
Body  subfusiform,  grub-shaped,  flattened  ventrally,  strongly  arched 
dorsally,  usually  curved  ventrally  (in  alcohol) .  Body  surface  covered 
with  low  globular  papillae  (to  naked  eye  surface  appears  almost 
smooth,  granular),  usually  with  thin  layer  of  sand  grains,  often  with 
an  almost  uniform  layer  of  larger  sand  grains  on  dorsum.  Prostomium 
and  buccal  segment  with  two  groups  of  tlu-ee  to  four  filiform  branchiae, 
pair  of  grooved  palps,  and  buccal  siphon  around  mouth.  First  seg- 
ment achaetous  (may  be  few  very,  delicate  capillary  setae),  second 
segment  with  two  small  neurosetae,  rest  of  segments  with  neuropodia 
as  slightly  elevated  lobes  and  about  four  (2-9)  amber-colored  nem-o- 
setae  with  curved  tips.  Without  distinct  notopodia ;  notosetae  poorly 
developed,  one  or  two  or  absent,  extremely  capillary,  easily  overlooked. 
Nephridial  papillae  on  ventral  side  between  segments  4  and  5  (setigers  3 
and  4,  since  the  first  segment  is  usually  achaetous).  Color:  In  life: 
Orange  yellow.     In  alcohol :  Light  gray  to  ashy. 

Remarks. — B.  granulata  Malmgren  is  herein  referred  to  B.  inhabilis, 
following  St0p-Bowitz  (1948a).  B.  sublaevis  Stimpson  should  also  be 
referred  here  and  not  to  B.  rugosa  (Hansen)  as  indicated  by  St0p- 
Bowitz  (1948a,  p.  37). 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  up  to  12.1 
miles  from  shore,  20-123.5  fms.,  on  bottoms  of  rocks,  stones,  gravel, 
and  mass  of  worm  tubes   (5  stations,  9  specimens).     Northwest 


MARINE    POLYCHAETE    WORMS — PETTIBONE  293 

Greenland:  1  mile  northwest  Conical  Rock,  25-60  fms.,  Bartlett, 
1938.  Bering  Sea:  Albatross  Sta.  3223,  54°26'  N.,  165°32'  W., 
56  fms.,  1890;  Bering  Strait,  13  fms.,  Dall,  1880.  Alaska:  Iliuliuk 
Harbor,  Unalaska,  1871,  and  New  Harbor,  Unga  Island,  under  stones, 
1872,  Dall;  Kodiak,  W.  J.  Fisher.  East  Coast  North  America: 
Off  Labrador,  95  fms..  Blue  Dolphin 'Eixpedition,  1949;  Bay  of  Fmidy, 
Maine,  48-90  fms.,  U.  S.  Fish  Commission. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian,  Alaskan, 
and  Canadian  Arctic,  Davis  Strait,  Greenland,  Novaya  Zemlya,Kara 
Sea.  Also  Iceland,  Faroes,  Norway  to  Danish  waters;  Labrador  to 
Maine;  Bering  Sea  to  Gulf  of  Alaska;  north  Japan  Sea.  In  low  water 
to  609  fathoms. 

Family  Scalibregmidae 

Body  inflated  anteriorly  or  short,  fusiform.  Integument  roughened 
or  tesseiated  and  divided  into  superficial  rings.  Prostomium  small, 
bilobed,  or  entire  and  with  frontal  horns;  two  ciliated  evaginable 
nuchal  grooves  on  each  side  of  prostomium  (fig.  33,  i).  Buccal 
segment  achaetous.  Parapodia  biramous,  rami  small,  with  or  without 
dorsal  and  ventral  cirri.  Setae  simple,  capillary,  some  single,  some 
forked.  Branchiae  limited  to  anterior  few  segments  (may  be  absent). 
Pygidium  with  or  without  anal  cirri.  Proboscis  eversible,  unarmed, 
globular,  smooth. 

Represented  by  a  single  species  from  Point  Barrow. 

Genus  Scalibregma  Rathke,  1843 
Scalihregma  inflatum  Rathke,  1843 

FiGUEE  33,  i-k 

Scalibregma  inflatum  Rathke,  1843,  p.  184,  pi.  9,  figs.  15-21. — Moore,  1923, 
p.  217.— Fauvel,  1927,  p.  123,  fig.  44,  a-f ;  1932,  p.  187;  1934a,  p.  50.— 
Augener,  1928,  p.  754.— Monro,  1930,  p.  163.— Annenkova,  1934,  p.  322; 
1937,  p.  176;  1938,  p.  186.— Okuda,  1938b,  p.  99.— Hartman,  1944a,  pp. 
336,  341;  1948,  p.  40.— St0p-Bowitz,  1946,  p.  67,  fig.  2;  1948b,  p.  24,  fig.  8.— 
Zatsepin,  1948,  p.  136,  pi.  33,  fig.  10.— Wesenberg-Lund,  1950a,  p.  36; 
1950b,  p.  86;  1951,  p.  81.— Berkeley  and  Berkeley,  1952,  p.  58,  figs.  119-121. 

Scalibregma  brevicauda  Verrill,  1873,  pp.  416,  422,  605;  1881,  p.  302,  pi.  9,  fig.  5. 

Scalibregma  minutum  Webster  and  Benedict,  1887,  p.  727. 

Description. — ^Length  up  to  100  mm.,  width  13  mm.,  45-60  segments 
(two  specimens  washed  ashore  after  a  storm  were  longer  than  any 
previously  recorded).  Body  divided  into  two  more  or  less  distinct 
regions,  anterior  inflated  portion  (anterior  15-17  setigers)  tapered 
anteriorly,  without  dorsal  and  ventral  cirri,  and  narrower  attenuated 
posterior  region  with  dorsal  and  ventral  cirri.     A  more  or  less  devel- 


294  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  103 

oped  ventral  groove  along  length  of  body,  better  developed  in  posterior 
region.  Integument  tesselated,  with  four  annuli  per  segment.  Pro- 
stomium  subquadrangular,  wider  anteriorly,  with  frontal  horns  on 
anterolateral  borders,  with  pair  of  evaginable  nuchal  organs  lateral 
to  prostomium  (when  inverted,  appear  as  diagonal  slits).  Buccal 
segment  achaetous,  forming  rugose  ring  around  prostomium.  Mouth 
ventral,  at  level  of  first  setigerous  lobes.  Proboscis  eversible,  globular, 
smooth,  unarmed.  Branchiae  four  pairs,  short,  tufted,  arborescent, 
inserted  posterior  to  notopodia  on  setigers  2-5,  increasing  in  size 
posteriorly.  Noto-  and  neuropodial  lobes  of  anterior  segments  small, 
projecting,  subequal.  Beginning  on  setigers  16-18,  dorsal  and  ventral 
cirri  develop  gradually,  becoming  elongate-conical,  flattened  or 
inflated.  Setae  silky,  iridescent,  capillary  (may  be  extra  long  in 
epitokous  pelagic  form).  Lateral  ciliated  organs  between  rami. 
Pygidium  with  five^o  seven,  short  to  long,  deciduous,  filiform  anal 
cirri  on  ventral  side.  Epitokous  sexual  forms  may  appear  in  surface 
waters,  massed  with  sexual  products,  with  extra  long  setae  and 
usually  with  highly  pigmented  dorsal  and  ventral  cirri.  Color: 
In  life :  Orange  yellow.  In  alcohol :  Without  color,  orange  yellow,  or 
sulfur  yellow  with  darker  pigmented  masses  in  the  outer  portions  of 
the  dorsal  and  ventral  cirri. 

Remarks. — Scalibregma  brevicauda  Verrill  is  herein  referred  to  S. 
injlatum.  Although  the  types  were  not  examined,  specimens  from 
the  New  England  region  identified  by  Verrill  as  S.  brevicauda  were 
observed.  The  types  of  S.  minutum  Webster  and  Benedict  from 
Eastport,  Maine,  were  examined  and  found  to  agree  with  S.  injlatum. 

New  records. — Arctic  Alaska:  Point  Barrow  base,  washed  ashore; 
off  Point  Barrow  base,  22.5-78  fms,,  up  to  15  miles  from  shore,  on 
bottoms  of  mud  and  various  combinations  of  mud,  sand,  gravel, 
stones,  rocks,  with  shells  and  worm  tubes  (9  stations,  17  specimens). 
West  Coast  North  America:  Washington  and  Puget  Sounds,  25 
fms.,  mud,  Pettibotie.  East  Coast  North  America:  Off  Labrador, 
17-35  fms..  Blue  Dolphin  Expeditions,  1950,  1951;  off  Nova  Scotia, 
Maine,  Massachusetts,  Rhode  Island,  Connecticut,  Long  Island 
Sound,  5-95.5  fms.,  U.  S.  Fish  Commission;  Cuttyhunk,  Massachu- 
setts, low  water,  in  sand,  Pettibone,  1950. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian,  Alaskan, 
and  Canadian  Arctic,  Davis  Strait,  Greenland,  Jan  Mayen,  Spits- 
bergen, Franz  Josef  Land,  Novaya  Zemlya,  Kara  Sea.  Also  Iceland, 
Faroes,  Norway  to  France,  Adriatic,  Gulf  of  Oman;  Labrador  to 
Long  Island  Sound;  Bering  Sea  to  central  California;  north  Japan 
Sea  to  Japan.  Southern  latitudes:  New  Zealand,  Magellan  Straits, 
South  Georgia,  Prince  Edward  Islands,  Kerguelen.  In  low  water  to 
1,333  fathoms;  epitokes  at  surface. 


MARESTE    POLYCHAETE    WORMS — ^PETTIBONE  295 

Family  Opheliidae 

Body  relatively  short,  often  grublike  or  lancet-shaped.  Integument 
smooth  or  granulated;  segmental  grooves  weak  or  obscured  by  super- 
ficial annuli.  Prostomium  small,  conical,  without  appendages;  pair 
of  eversible  nuchal  organs  (fig.  33,  I,  o).  Proboscis  eversible,  saclike, 
unarmed.  Parapodia  biramous,  lobes  reduced  or  lacking;  setae  simple, 
capillary  (fig.  33,  m,  q).  Without  dorsal  cirri;  rarely  with  ventral 
cirri.  Branchiae  paired,  cirriform,  inserted  near  parapodia,  extending 
along  most  of  body  (may  be  lacking).  Pygidium  forming  an  anal 
cylinder,  a  flap,  or  knob,  with  or  without  cirri  (fig.  33,  n,  p). 

Represented  by  two  genera  and  two  species  from  Point  Barrow. 
Both  genera  have  paired  cirriform  branchiae  along  most  of  length  of 
body;  without  lateral  eyes  along  body. 

Key  to  the  genera  of  Opheliidae  from  Point  Barrow 

1.  Body  lancet-shaped;  ventral  surface  thick,  solelike,  formed  by  2  thick  longi- 
tudinal muscular  columns,  with  midventral  and  lateral  grooves  along  length 
of  body.     Parapodial  lobes  small,  along  lateral  grooves  (fig.  33,  m). 

Ammotrypane  (p.  295) 
Body  cylindrical,  grub-shaped,  without  midventral  or  lateral  grooves.     Para- 
podial lobes  absent,  the  simple,  capillary  noto-  and  neurosetae  emerging 
directly  from  body  wall  (fig.  33,  q) Travisia  (p.  296) 

G^nus  Ammotrypane  Ratlike,  1843 
Ammotrypane  breviata  Ehlers,  1913 

Figure  33,  l-n 

Ammotrypane  breviata  Ehlers,  1913,  p.  523,  pi.  39,  figs.  1-7. — Monro,  1930,  p. 

165;  1936,  p.  165. 
Ophelina  groenlandica  St0p-Bowitz,  1948b,  p.  20,  fig.  6. 

Description. — Length  6-7  mm.,  width  0.3  mm.,  setigerous  segments 
26  or  27  (up  to  34  mm.  long,  2  mm.  wide,  26-28  segments — Monro, 
1930,  1936).  Body  rigid,  tapered  at  both  ends,  smooth,  without 
superficial  segmental  divisions,  Prostomium  conical,  the  anterior  tip 
constricted  to  form  a  small  globular  palpode;  prostomial  tip  may  be 
partly  telescoped  into  rest  of  prostomium;  pair  of  eversible  nuchal 
organs  at  level  of  parapodia.  Parapodia  with  two  groups  of  fine 
capillary  setae,  the  dorsal  group  slightly  longer  than  the  ventral. 
Branchiae  begin  on  setiger  2,  absent  on  last  three  to  five  setigers; 
branchiae  slender,  cirriform,  subequal.  Pygidium  forming  a  short, 
somewhat  contractile  anal  cylinder,  closed  on  aU  sides,  somewhat 
lobulate  or  papillate  on  open  posterior  end.  Eversible  proboscis 
saclike,  smooth.  Color:  In  alcohol:  Without  color  or  may  be  darkly 
pigmented  laterally  along  posterior  three  setigerous  segments  and 
proximal  part  of  anal  tube. 


296  PROCEEDINGS    OE   THE    NATIONAL   MUSEUM  vol.  103 

Remarks. — The  two  small  specimens  from  Point  Barrow  (6-7  mm. 
long)  follow  closely  the  description  of  St0p-Bowitz  for  Ophelina  groen- 
landica,  including  size.  They  seem  to  agree  with  the  description  by 
Ehlers  of  A.  breviata  from  the  Antarctic,  although  the  Antarctic 
specimens  attain  a  much  larger  size  (29  mm.  long). 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  up  to  7.5 
miles  from  shore,  29-36  fms.,  on  bottoms  of  gravel,  stones,  large 
perforated  rocks  (2  stations,  2  specimens). 

Distribution. — ^Arctic  Alaska,  East  Greenland.  Southern  latitudes: 
South  Orkney  and  Falkland  Islands,  Kaiser- WUhelm  II  Land.  In 
10-214  fathoms. 

Genus  Travisia  Johnston,  1840 
Travisia  camea  Verrill,  1873 

Figure  33,  o-q 

Travisia  cornea  Verrill,  1873,  pp.  431,  434,  508,  604;  1881,  p.  302,  pi.  8,  fig.  1.— 
Sumner,  Osburn,  and  Cole,  1913,  p.  629.— Hartman,  1944a,  pp.  336,  341,  pi. 
19,  fig.  6  (as  probably  Ophelia  limacina). 

Travisia  forbesi  Murdoch,  1885,  p.  154  (not  T.  forbesii  Johnston,  1840). 

Description. — Length  up  to  59  mm.,  width  8  mm.,  segments  25-29 
(specimens  washed  ashore  at  Point  Barrow  much  larger  than  any 
previously  recorded).  Body  rounded,  cylindrical,  grub-shaped, 
tapered  toward  each  end.  Segments  of  posterior  third  may  be  some- 
what imbricated  or  telescoped;  triannulate  in  middle  region.  Integu- 
ment with  granulations  or  beading  rather  uniform  (not  larger  on 
posterior  part  of  segment  as  in  T.  pupa  Moore).  Prostomium  small, 
conical,  acutely  to  bluntly  pointed,  without  eyes.  Branchiae  begin  on 
setiger  2  (rarely  on  setiger  3),  continuing  posteriorly,  absent  on  last 
four  or  five  segments,  cirriform,  inserted  posterior  to  notosetae. 
Without  parapodial  lobes,  the  small,  slender,  capillary  noto-  and  neuro- 
setae  emerging  directly  from  body  wall.  In  posterior  third  of  body, 
area  in  region  of  setae  may  be  somewhat  inflated  but  without  enlarged 
lateral  lobes  extending  posteriorly  (as  in  T.  forbesii  Johnston  and 


Figure  33. — Cirratulidae:  a,  Cirratulus  cirratus,  dorsal  view  anterior  end;  h,  same,  para- 
podium  from  anterior  region;  c,  same,  acicular  crotchet;  d,  Chaetozone  setosa,  dorsal 
view  anterior  end.  Flabelligeridae:  e,  Flabelligera  affinis,  parapodium;  /,  same,  hooked 
pseudoconipound  neuroseta;  g,  same,  pedunculate  papillae — (1)  from  tip  of  notopodium, 
(2)  on  body;  k,  Brada  inhahilis,  ventral  view  anterior  end.  Scalibregmidae:  i,  Scalihregma 
inflatum,  dorsal  view  anterior  end;  /,  same,  parapodium  from  middle  region;  k,  same, 
parapodium  from  posterior  region.  Opheliidae:  /,  Ammotrypane  breviata,  ventral  view 
anterior  end,  proboscis  extended;  w,  same,  lateral  view  middle  part  of  body;  n,  same, 
ventral  view  posterior  end;  o,  Travisia  camea,  dorsal  view  anterior  end;  p,  same,  dorsal 
view  posterior  end;  q,  same,  parapodium.  Capitellidae:  r,  Capitella  capitata,  dorsal  view 
anterior  end;  /,  same,  dorsal  view  posterior  part  thoracic  region;  t,  same,  parapodium 
from  abdominal  region;  u,  same,  hooded  crotchet.  (For  explanation  of  symbols,  see 
p.  210.) 


MARINE    POLYCHAETE    WORMS — PETTIBONE 


297 


P     Q 

Figure  33. — For  explanation  see  facing  page. 


298  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  103 

T.  brevis  Moore).  With  lateral  sensory  pits  between  rami  appearing 
as  small  oval  slits.  Nephridial  pores  ventral  to  nem-opodia  on  setigers 
3-14.  Pygidium  small,  cylindrical,  longitudinally  fmrowed.  Color: 
In  life:  Flesh.    In  alcohol:  Whitish. 

New  records. — Arctic  Alaska:  Point  Barrow  base,  washed  ashore, 
October  16,  17,  1949;  August  10,  1950  (12  specimens).  East  Coast 
North  America:  Off  Massachusetts,  Long  Island  Sound,  3-19  fms., 
soft  mud,  U.  S,  Fish  Commission;  Woods  Hole  region,  Massachusetts, 
Pettibone,  1950. 

Distribution. — Arctic  Alaska.  Massachusetts  to  Long  Island  Sound. 
In  3-19  fathoms. 

Family  Capitellidae 

Body  cylindrical,  wealdy  divided  into  2  regions:  Anterior  thoracic 
region  somewhat  inflated,  often  reticulated,  parapodial  lobes  rudi- 
mentary; posterior  abdominal  region  longer  and  thinner,  with  rows 
of  crotchets  on  somewhat  inflated  tori.  Prostomium  conical,  more 
or  less  retractile,  without  appendages,  with  or  without  eyes,  with 
pair  of  eversible  nuchal  organs.  Buccal  segment  achaetous  (may  be 
fused  with  prostomium  as  in  Capitella).  Parapodia  biramous,  re- 
duced to  bundles  of  capillary  setae  or  to  dorsal  and  ventral  tori 
bearing  row  of  crotchets  which  have  long  manubrium  and  recurved 
hooded  rostrum  (fig.  33,  r-^).  Without  dorsal,  ventral,  or  anal  cirri. 
With  or  without  branchiae.  Proboscis  eversible,  globular,  unarmed, 
papillate.     Constructs  tubes. 

Represented  by  a  single  species  from  Point  Barrow. 

Genus  Capitella  Blainville,  1828 
Capitella  capitata  (Fabricius,  1780) 

Figure  33,  r-u 

Lumbricus  capitatus  Fabricius,  1780,  p.  279. 

Capitella  capitata  Webster  and  Benedict,  1884,  p.  730;  1887,  p.  744. — Ehlers, 
1913,  p.  543.— Eliason,  1920,  p.  63.— Fauvel,  1927,  p.  154,  fig.  55,  a-h.— 
Augener,  1928,  p.  749.— Monro,  1930,  p.  163.— Anneukova,  1934,  p.  322; 
1937,  p.  179;  1938,  p.  189.— Hartman,  1942a,  p.  69;  1944a,  pp.  334,  341, 
pi.  34,  fig.  3;  1945,  p.  37;  1947,  p.  404,  pi.  43,  figs.  1,  2;  1948,  p.  41;  1951, 
p.  101.— Berkeley  and  Berkeley,  1943,  p.  129;  1952,  p.  100,  figs.  206-208.— 
Gorbunov,  1946,  p.  39.— Thorson,  1946,  p.  110,  fig.  60.— Zatsepin,  1948, 
p.  138,  pi.  33,  fig.  14. — Hartman  and  Reish,  1950,  p.  40. — Wesenberg-Lund, 
1950a,  p.  39;  1950b,  p.  91;  1951,  p.  84. 

Capitella  capitata  var.  antarctica  Monro,  1930,  p.  164. 

Description. — Length  23  mm.,  width  1  mm.  (up  to  120  mm. — 
Augener,  1928).  Body  attenuated  slightly  anteriorly,  much  more 
so  posteriorly,  fragile.  Prostomium  and  achaetous  buccal  segment 
fused,  forming  wide  obtuse  cone,  somewhat  flattened  dorsoventrally, 


MARINE    POLYCHAETE    WORMS — PETTIBONE  299 

usually  concave  dorsally/witli''pair^of  small  ventral  eyes  and  pair  of 
crescent-shaped  nuchal  slits  behind  eyes  (best  seen  from  lateral  view) , 
Thoracic  region  smooth,  without  reticulated  integument,  consisting 
of  first  9  setigerous  segments,  each  of  first  7  with  four  bundles  of 
capUlary  setae  which  on  setiger  7  may  be  only  capillaries,  only  crotch- 
ets, or  both  (in  young  specimens,  crotchets  may  be  present  from 
setiger  4) .  Setigers  8  and  9  modified,  having  neuropodia  with  row  of 
hooded  crotchets,  and  with  large,  recurved,  amber-colored  acicular 
genital  hooks  located  middorsally  between  these  two  segments. 
Abdominal  segments  with  slightly  raised  dorsal  and  ventral  para- 
podial  tori  each  bearing  a  row  of  hooded  crotchets.  Without  spe- 
cialized branchiae  or  lateral  organs.  Pygidium  a  short  ring,  may  be 
inflated.  Color:  In  alcohol:  Without  color  or  reddish  brown. 
Females  with  large  yolky  eggs  (Point  Barrow,  August  9,  30,  1949; 
September  1,  6,  1949.) 

Remarks. — According  to  Fauvel  and  others,  in  C.  capitata  the  genital 
hooks  on  setigers  8  and  9  are  present  only  in  the  males,  the  females 
having  only  crotchets.  On  all  the  specimens  from  Point  Barrow, 
genital  hooks  are  present,  including  some  females  containing  large 
yolky  eggs.  They  agree  in  all  other  respects  with  C.  capitata.  Having 
genital  hooks  in  both  males  and  females,  they  would  fall  under  the 
very  closely  related  Capitellides  Mesnil.  They  differ  from  C.  giardi 
Mesnil  in  the  larger  number  of  genital  hooks.  They  are  herein  re- 
ferred to  the  cosmopolitan  C.  capitata  with  the  tentative  idea  that  the 
relative  development  of  genital  hooks  in  the  female  may  prove  to  be 
a  variable  character. 

New  records. — Arctic  Alaska:  Eluitkak  Pass,  Elson  Lagoon  near 
Point  Barrow,  6.6  fms.;  off  Point  Barrow  base,  up  to  7.5  miles  from 
shore,  23.3-70  fms.,  on  bottoms  of  stones  and  various  combinations 
of  gravel,  stones,  rocks,  large  perforated  rocks  (8  stations,  26  speci- 
mens). Bering  Sea:  St.  Paul  Island,  Pribilofs,  in  holdfasts  with 
Fabricia  sabella  and  lumbrinerids,  W.  L.  Hahn,  1911.  West  Coast 
North  America:  Corona  del  Mar,  California,  from  branchial  cavity 
of  crab,  Loxorhynchus  grandis,  MacGinitie,  1948.  East  Coast  North 
America:  Off  Labrador,  6  fms.,  mud-sand.  Blue  Dolphin  Expedition, 
1949;  off  Massachusetts,  15  fms.,  U.  S.  Fish  Commission;  Woods  Hole 
region,  Massachusetts,  Pettibone,  1951. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian,  Alaskan, 
and  Canadian  Arctic,  Davis  Strait,  Greenland,  Spitsbergen,  Novaya 
Zemlya,  Kara  Sea.  Also  Iceland,  Faroes,  Norway  to  France,  Madeira, 
Mediterranean,  Black  Sea;  Hudson  Bay  to  North  Carolina,  Texas; 
Bering  Sea  to  southern  California;  north  Japan  Sea.  Southern  lati- 
tudes :  Magellan  Straits,  South  Georgia,  Bouvet  Island,  South  Africa, 
Kerguelen.     In  low  water  to  500  fathoms. 

261112—54 7 


300  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  103 

Family  Arenicolidae 

Body  elongate,  cylindrical,  integument  reticulated  and  divided  into 
secondary  annuli.  Body  divided  into  two  or  three  distinct  regions: 
(1)  anterior,  abranchial;  (2)  abdominal,  branchial,  each  segment  with 
pair  of  dorsally-borne  branched  gills;  (3)  caudal,  achaetous  and 
abranchial;  this  region  may  be  lacking.  Prostomium  small,  simple 
or  trilobed,  without  appendages;  with  nuchal  groove  (fig.  34,  a). 
Parapodia  biramous;  notopodia  conical,  setigerous,  with  spiny,  capU- 
lary  setae;  neuropodia  with  sigmoid  crotchets  in  semicircular  tori. 
Without  dorsal,  ventral,  and  anal  cirri.  Anus  terminal.  Proboscis 
eversible,  globular,  papillate,  unarmed.     Constructs  tubes  or  burrows. 

Represented  by  a  single  species  from  Point  Barrow. 

Genus  Arenicola  Lamark,  1801 
Arenicola  glacialis  Murdoch,  1884 

Figure  34,  a 

Arenicola  glacialis  Murdoch,  1884,  p.  522;  1885,  p.  155. — Ashworth,  1910,  p.  24, 
figs.  10-14;  1912,  p.  Ill,  figs.  46,  47,  pi.  6;  1924,  p.  3,  fig.  1. 

Description. — Length  up  to  205  mm.,  width  11  mm.,  17  setigerous 
segments.  Body  cylindrical,  tapering  slightly  anteriorly  and  gradu- 
ally posteriorly.  Prostomium  small,  may  be  pretty  well  retracted  in 
nuchal  groove;  median  lobe  small,  lateral  lobes  larger,  curved  (fig. 
34,  a).  Body  divided  into  three  regions:  (1)  anterior  region  of  six 
abranchial  setigers,  notopodia  conical  with  spiny  capillary  setae, 
neuropodia  low  tori  with  few  sigmoid  crotchets;  (2)  branchial  region 
of  11  setigers  each  divided  into  five  superficial  annuli,  with  short, 
bushy,  arborescent  branchiae  at  base  of  conical  notopodia,  with  gill 
axes  short,  branching  one  or  two  times  or  simple;  neuropodia  raised 
tori  with  long  row  of  crotchets;  on  more  posterior  segments,  tori 
approach  midventral  line;  (3)  caudal  achaetous  and  abranchial  region 
with  papillae  feebly  developed,  without  processes  (as  in  A.  cristata). 
Nephridial  pores  six  pairs,  on  setigers  4-9,  posterior  to  upper  part  of 
neuropodial  tori.     Pair  of  statocysts  on  peristomium,  open  to  exterior. 


Figure  34. — Arenicolidae:  a,  Arenicola  glacialis,  prostomium.  Maldanidae:  b,  Praxillella 
praeUrmissa,  ventral  view  anterior  end;  c,  same,  frontal  view  cephalic  plate;  d,  same, 
pygidial  funnel;  g,  same,  posterior  end;  /,  same,  crotchet  from  neuropodium;  g,  Maldane 
sarsi,  lateral  view  anterior  end;  h,  same,  lateral  view  posterior  end;  i,  Nicomache  lum- 
bricalis,  lateral  view  anterior  end;  /,  same,  lateral  view  posterior  end;  k,  Nicomache 
per  sonata,  lateral  view  posterior  end;  /,  Petaloproctus  tenuis,  lateral  view  anterior 
end;  m,  same,  lateral  view  posterior  end.  Sabellariidae:  n,  Idanthyrsus  armatus,  dorsal 
view  anterior  end;  o,  same,  lateral  view  anterior  end;  p,  same,  palea  from  inner  row; 
q,  same,  palea  from  outer  row.     (For  explanation  of  symbols,  see  p.  210.) 


MARINE    POLYCHAETE    WORMS — PETTIBONE  301 

■  set  19, 


Figure  34. — For  explanation  see  facing  page. 


302  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  103 

with  numerous  sand  grains  as  statoliths.  Color  :  In  life :  Deep  orange 
to  brownish  red.     In  alcohol:  Bluish  to  grayish  black  or  brown. 

New  records. — Arctic  Alaska:  Point  Barrow  base,  washed  ashore 
August  21,  24,  28,  September  6,  12,  19,  22,  October  16,  1949  (54 
specimens);  off  Point  Barrow  base,  along  shore,  1.7  fms.,  on  bottom 
of  gravel  with  mud  (2  stations,  3  specimens) . 

Distribution. — Alaskan  and  Canadian  Arctic.     In  1.7-3  fathoms. 

Family  Maldanidae 

Body  cylindrical,  truncated  at  both  ends,  without  differentiated 
regions,  segments  usually  few,  some  much  longer  than  wide.  Pro- 
stomium  a  small  cephalic  crest  fused  with  achaetous  buccal  segment, 
forming  ovoid  head,  without  appendages,  with  or  without  oblique 
limbate  brim  or  plate  (fig.  34,  b,  i,  I) ;  pair  of  nuchal  slits  on  either  side 
cephalic  crest.  In  anterior  region  with  numerous  glandular  cells, 
diffused  or  grouped  in  clear  bands  alternating  with  pigmented  rings. 
Parapodia  biramous;  notopodia  with  capillary  setae;  neuropodia  with 
projecting  tori  bearing  rows  of  crotchets  (fig.  34,/)  with  dentate  ros- 
trum and  rostral  hairs;  the  crotchets  may  be  replaced  by  few  stout 
acicular  spines  on  first  few  setigers.  Without  dorsal  or  ventral  cirri; 
usually  without  branchiae.  Pygidium  a  limbate  plate,  spatulate  or 
funneUike  (fig.  34,  h,  j,  m).  Proboscis  eversible,  unarmed,  globular. 
Live  in  tubes  in  sand  or  mud;  tube  cylindrical,  membranous,  covered 
with  thin  to  thick  layer  of  mud,  sand,  gravel,  more  or  less  resistant. 

Represented  by  four  genera  and  five  species  from  Point  Barrow.  All 
the  genera  are  without  specialized  collarettes,  without  special  glandular 
ventral  shield  on  setiger  8,  with  neuropodial  crotchets  in  single  rows. 

Key  to  the  genera  of  Maldanidae  from  Point  Barrow 

1.  Head  with  limbate  plate  (fig.  34,  h,  g) 2 

Head  without  limbate  plate  (fig.  34,  i,  I).  With  1-2  stout  acicular  setae  in 
neuropodia  of  first  3  setigers.  Pygidium  funnel-shaped  or  forming  concave 
plate,  with  or  without  cirri;  anus  opening  on  surface  of  plate 3 

2.  Head  with  low  cephalic  crest,  bordered  by  oblique  limbate  plate,  entire  or 

notched  laterally  (fig.  34,  h,  c) .    With  ventral  crotchets  beginning  on  setiger  1. 
Pygidium  funnel-shaped,  bordered  with  cirri,  with  anus  in  center  of  somewhat 

protruding  anal  cone  within  the  funnel  (fig.  34,  d) Praxillella  (p.  303) 

Head  with  middorsal  convex  cephalic  crest  or  keel,  bordered  by  limbate  plate, 
divided  into  3  parts  by  2  deep  lateral  grooves  (fig.  34,  g).  Without  neuro- 
setae  on  setiger  1 ;  crotchets  begin  on  setiger  2.  Pygidium  a  slightly  oblique 
oval  plate,  bordered  by  rim  which  is  incised  laterally,  smooth  or  denticled; 
anus  dorsal  to  pygidial  disc  (fig.  34,  h) Maldane  (p.  303) 

3.  Pygidium  sHghtly  obhque,  funnel-shaped,  bordered  with  cirri  (fig.  34,  j,  k). 

Nicomache  (p.  304) 

Pygidium  an  asj'^mmetrical,  foliaceous,  concave  plate,  reduced  dorsally,  wider 

ventrally,  with  border  smooth  or  crenulate  (fig.  34,  m)  .Petaloproctus  (p.  306) 


MARINE    POLYCHAETE    WORMS — PETTIBONE  303 

Genus  Praxillella  Verrill,  1881 
Praxillella  praetertnissa  (Malmgren,  1865) 

Figure  34,  b-f 

Praxilla  praetermissa  Malmgren,  1865,  p.  191;  1867,  p.  100,  pi.  11,  fig.  62. 
Praxillella  praetermissa  Verrill,    1881,   pp.   298,   305,   309,   312.— Webster  and 

Benedict,  1887,  p.  746.— Arwidsson,  1907,  p.  192,  figs.— Eliason,  1920,  p. 

66.— Augener,    1928,   p.   762.— Annenkova,    1937,  p.   182;   1938,  p.    194.— 

Zatsepin,  1948,  p.  142,  pi.  35,  fig.  1.— Wesenberg-Lund,  1948,  p.  41,  figs. 

20,  21;  1950a,  p.  43;  1950b,  p.  98;  1951,  p.  93. 
Clymene  {Praxillella)  praetermissa  Fauvel,  1927,  p.  179,  fig.  62,  a-e. 

Description. — Length  45  mm.,  width  1  mm,  (up  to  120  mm.  long, 
2.5  mm.  wide — Malmgren,  1865),  19  setigerous  segments,  4-5  achaetous 
segments  with  rudimentary  parapodia.  Head  with  oblique  limbate 
plate,  with  border  entire,  slightly  or  deeply  incised  laterally,  slightly 
incised  posteriorly,  terminating  anteroventraUy  in  obtuse  cone  (not 
produced  into  digitiform  process);  median  cephalic  crest  long;  nuchal 
grooves  one-half  to  two-thirds  its  length.  Neuropodia  of  anterior 
three  setigers  with  2-10  slightly  modified  crotchets.  Pygidial  funnel 
with  flat  floor,  with  projecting  anal  cone  with  larger  valvule  on  ventral 
side  of  cone;  border  of  funnel  with  20-38  subequal  papillae  except 
for  a  longer  ventral  one.  Proboscis  papillate  basally.  Color:  In 
life:  Yellow  with  wide,  red  and  white  bands  on  some  anterior  segments. 
In  alcohol:  Brownish  to  gray,  with  whitish  bands  anterior  to  setae 
on  setigers  4-8.  Tube  :  Of  clay,  sand,  small  bits  of  rock;  may  be  quite 
resistant. 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  up  to  7.5 
miles  from  shore,  20-36  fms.,  on  various  combinations  of  mud,  sand, 
gravel,  stones,  rocks,  large  perforated  rocks,  shells  (7  stations,  10 
specimens).  East  Coast  North  America:  Off  Labrador,  5-8  fms., 
Blv£  Dolphin  Expedition,  1949;  off  Cape  Cod,  Massachusetts,  14-19 
fms.,  U.  S.  Fish  Commission. 

Distribution. — Widely  distributed  in  the  Arctic:  Alaskan  Arctic, 
Davis  Strait,  Greenland,  Spitsbergen,  White  Sea,  Novaya  Zemlya, 
Kara  Sea.  Also  Iceland,  Faroes  to  English  Channel,  Mediterranean; 
Labrador  to  Massachusetts;  north  Japan  Sea.     In  7-1,111  fathoms. 

Genus  Maldane  Grube,  1860 
Maldane  sarsi  Malmgren,  1865 

Figure  34,  g,  h 

Maldane  sarsi  Malmgren,  1865,  p.  188;  1867,  p.  99,  pi.  10,  fig.  57. — Arwidsson, 
1907,  p.  251,  figs.— Eliason,  1920,  p.  66.— Fauvel,  1927,  p.  197,  fig.  69,  a-i; 
1932,  p.  202.— Augener,  1928,  p.  759.— Annenkova,  1937,  p.  179;  1938,  p. 
192.— Okuda,  1939,  p.  239.— Berkeley  and  Berkeley,  1942,  p.  199.— Hartman, 


304  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  103 

1944a,  pp.  335,  342,  pi.  33,  figs.  3,  4;  1951,  p.  106.— Zatsepin,  1948,  p.  142, 

pi.  33,  fig.  3.— Wesenberg-Lund,  1948,  p.  48,  figs.  24,  25;  1950a,  p.  44;  1950b, 

p.  100;  1951,  p.  94. — Hartman  and  Reish,  1950,  p.  37. 
Maldane  sarsi  var,  antardica  Arwidsson,   1911a,  p.  32,  figs. — Munro,  1930,  p. 

169;  1936,  p.  168. 
Maldane  sarsi  var.  Moore,  1923,  p.  237. 
Maldane  sarsi  var.  tropica  Monro,  1937,  p.  307,  fig.  24. 

Description. — Length  up  to  110  mm.,  width  3.9  mm.,  setigerous 
segments  19,  two  antanal  achaetous  segments.  Head  with  long, 
convex  keel,  ending  in  blunt  process  above  mouth,  bordered  by  limbate 
plate,  with  border  smooth,  usually  incised  laterally,  rather  high  pos- 
teriorly; nuchal  organs  short,  divergent;  without  ocelli.  Anterior 
segments  biannulate.  With  notosetae  only  on  first  setiger;  neuro- 
podial  crotchets  begin  on  setiger  2.  Two  antanal  segments  short, 
with  achaetous  tori  slightly  marked.  Pygidium  an  oblique  flat,  oval 
plate,  with  border  usually  slightly  incised  laterally,  smooth  or  slightly 
crenulate  ventrally.  Anus  a  folded  cushion  dorsal  to  and  close  to 
pygidial  rim.  Tube:  Free,  circular,  thin  to  thick  walled,  of  fine 
agglomerated  mud  and  sand  grains. 

New  Records. — Arctic  Alaska:  Off  Point  Barrow  base,  3  miles 
from  shore,  20.3  fms.,  on  bottom  of  mud,  gravel,  stones,  shells  (1 
station,  1  specimen),  Bering  Sea:  Albatross  Sta.  3603,  55°23'  N., 
170°31'  W.,  1,771  fms.,  1895.  Canadian  Arctic:  Northwest  shore 
Daniels  Island,  mouth  Newell  Sound,  Frobisher  Bay,  Baffin  Land, 
10-30  fms.,  Bartlett.  East  Coast  North  America:  Off  Labrador, 
30  fms..  Blue  Dolphin  Expedition,  1950;  off  Nova  Scotia,  Maine, 
Massachusetts,  Rhode  Island,  14-487  fms.,  U.  S.  Fish  Commission., 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian,  Alaskan 
and  Canadian  Arctic,  Davis  Strait,  Greenland,  Spitsbergen,  Novaya 
Zemlya,  Kara  Sea.  Also  Iceland,  Norway  to  France;  Labrador  to 
Rhode  Island,  Louisiana,  Mississippi,  eastern  Texas;  Bering  Sea  to 
southern  California;  north  Japan  Sea  to  Japan;  South  Arabian  coast 
(var.  tropica);  Bay  of  Bengal.  Southern  latitudes:  South  Georgia, 
South  Shetlands,  Palmer  Archipelago  (var.  antarctica) ;  south  Austraha. 
In  low  water  to  1,771  fathoms. 

Genus  Nicomache  Malmgren,  1865 

Both  species  have  buccal  segment  fused  with  prostomium  forming 
hood-shaped  head,  irregularly  rounded,  with  thickened  anteroventral 
border,  forming  projecting  upper  lip  (fig.  34,  i).  First  three  setigerous 
segments  each  with  one  or  two  stout,  acicular,  spinelike  nem"0setae. 
Some  of  notopodia  with  long,  hairlike  sinuous  dorsal  setae.  Pygidium 
with  anus  terminal  in  center  of  short,  slightly  oblique  funnel,  with 
papillae  on  border  of  funnel  (fig.  34,  j,  k) . 


MARINE    POLYCHAETE    WORMS — PETTIBONE  305 

Key  to  the  species  of  Nicomache  from  Point  Barrow 

1.  Single  achaetous  antanal  segment  (fig.  34,  k) N.  personata 

Two  achaetous  antanal  segments  (fig.  34,  j) N.  lumbricalis 

Nicomache  lumbricalis  (Fabricius,  1780) 

Figure  34,  i,  j 

Sabella  lumbricalis  Fabricius,  1780,  p.  374. 

Nicomache  lumbricalis  Malmgren,  1867,  p.  99,  pi.  10,  fig.  60. — Webster  and 
Benedict,  1884,  p.  731;  1887,  p.  745.— Arwidsson,  1907,  p.  86,  pi.  8,  figs. 
244,  245.— Fauvel,  1927,  p.  190,  fig.  66,  a-i.— Augener,  1928,  p.  764.— Monro, 
1930,  p.  173.— Annenkova,  1934,  p.  322;  1937,  p.  181;  1938,  p.  193.— Hartman, 
1944a,  pp.  335,  342;  1948,  p.  42.— Zatsepin,  1948,  p.  144,  pi.  26,  fig.  1.— 
Wesenberg-Lund,  1948,  p.  23,  figs.  10,  11;  1950a,  p.  41;  1950b,  p.  95;  1951, 
p.  90.— Berkeley  and  Berkeley,  1952,  p.  54,  figs.  Ill,  112. 

Nicomache  carinaia  Moore,  1906b,  p.  242,  pi.  11,  figs.  36-39,  pi.  12,  figs.  43,  44; 
1923,  p.  227. 

Nicomache  lumbricalis  var.  borealis  Arwidsson,  1907,  p.  94,  figs.;  1922,  p.  6. — 
Eliason,  1920,  p.  65. 

Description. — Length  40-60  mm.,  width  2.5-3  mm.,  setigerous 
segments  21-22,  two  short  achaetous  antanal  segments  (Point  Barrow 
and  Labrador  specimens  with  22  setigers;  type  of  N.  carinaia  with  21 ; 
up  to  160  mm.  long,  5  mm.  wide — Fauvel,  1927).  Prostomium  with- 
out ocelli;  nuchal  organs  S-shaped.  Pygidial  funnel  with  14-30 
subequal,  pointed,  triangular  papillae.  Color:  In  alcohol:  Reddish 
brown  on  sides  and  dorsal  surface  of  head  and  first  three  setigers. 
Tube:  Sandy,  with  small  pieces  of  rock,  with  smooth  membranous 
lining  (Point  Barrow),  or  thick,  sandy,  and  coiled  (Labrador;  small 
ones  on  terebeUid  tubes) . 

Remarks. — They  may  share  their  tubes  with  the  commensal  poly- 
noid  Enipo  gracilis  Verrill  (=E.  cirrata  TreadweU)  reported  by  Berke- 
ley and  Berkeley  from  Alaska;  also  from  off  Nova  Scotia  and  Mas- 
sachusetts (in  USNM). 

New  records. — Arctic  Alaska:  Off  Point  Barrov/,  up  to  12.1  miles 
from  shore,  24.7-123.5  fms.,  on  bottoms  of  mud,  worm  tubes,  stones, 
and  various  combinations  of  rocks,  stones,  gravel  (6  stations,  18 
specimens).  Bering  Sea:  Cape  Prince  of  Wales,  23  fms.,  Dall,  1874. 
Albatross  Sta.  3311,  53°  59'  N.,  166°  29'  W.,  85  fms.,  and  Sta.  3313, 
54°  or  N.,  166°  27'  W.,  68  fms.,  1890.  Alaska:  Albatross  Sta.  2848, 
Unalaska  to  Cook  Island,  1888;  Eastern  Harbor,  Sitka,  15  fms.,  Dall, 
1874.  Kamchatka:  Albatross  Sta.  3644,  51°  09'  N.,  157°  48'  W.,  96 
fms.,  1896.  East  Coast  North  America:  Off  Labrador,  30-125 
fms..  Blue  Dolphin  Expeditions,  1949,  1950,  1951;  off  Nova  Scotia, 
Bay  of  Fundy,  Maine,  New  Hampshire,  Massachusetts,  26-150  fms., 
U.  S.  Fish  Commission. 

Distribution. — Widely    distributed    in    the    Arctic:  Siberian    and 


306  PROCEEDINGS    OF   THE    NATIONAL   RIUSEUM  vol.  103 

Alaskan  Arctic,  Greenland,  Jan  Mayen,  Spitsbergen,  White  Sea, 
Novaya  Zemlya,  Kara  Sea,  Also  Bering  Sea  to  Lower  California; 
north  Japan  Sea;  Iceland,  Norway  to  English  Channel,  Santander; 
Labrador  to  Massachusetts;  South  Africa.  In  low  water  to  1,400 
fathoms. 

Nicomache  personata  Johnson,  1901 

Figure  34,  k 

Nicomache  personata  Johnson,   1901,  p.  419,  pi.  13,  figs.  134-139. — Arwidsson, 

1922,  p.  7,  pi.  1,  figs.  5,  6.— Hartman,  1948,  p.  41,  fig.  11,  d-g.— Berkeley 

and  Berkeley,  1952,  p.  54,  figs.  109,  110. 
Nicomache  minor  Arwidsson,  1907,  p.  100,  pi.  2,  figs.  68-73;  pi.  8,  figs.  252-256. — 

Augener,  1928,  p.  765.— Annenkova,  1937,  p.  181;  1938,  p.  193. 
Nicomache  maculata  Arwidsson,  1911b,  p.  209,  pi.  18,  figs.  13-19;  pi.  19,  figs. 

27-30.— Fauvel,  1927,  p.  191,  fig.  66,  k-r. 

Description. — Up  to  110  mm.  long,  2  mm.  wide,  22  or  23  setigerous 
segments,  single  achaetous  antanal  segment  (Point  Barrow  specimen 
with  23  setigers,  Puget  Sound  specimen  with  22).  Prostomium  with 
numerous  ocelli  (may  be  absent) ;  nuchal  clefts  short,  arched.  Pygidial 
funnel  with  16-26  papillae  which  are  short,  subequal,  triangular,  or 
very  unequal,  some  being  longer,  some  very  short,  few  may  be  bifid 
or  trifid.  Color:  In  alcohol:  Without  color,  or  brownish,  especially 
on  anterior  end.  Tube  :  Thick  walls  of  coarse  sand  particles  cemented 
together. 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  up  to  8 
miles  from  shore,  21.7-75.5  fms.,  on  various  combinations  of  pebbles, 
rocks,  gravel,  stones  (3  stations,  5  specimens).  Bering  Sea:  Alba- 
tross Sta.  3289,  56°  44'  N.,  159°  16'  W.,  16  fms.,  1890.  West  Coast 
North  America  :  Security  Bay,  Alaska,  Jones;  Washington  and  Puget 
Sounds,  low  water  to  61  fms.,  Pettiboiie. 

Distribution. — Arctic  Alaska,  Spitsbergen;  Bering  Sea  to  Washing- 
ton; north  Japan  Sea;  Scandinavia,  Ireland,  North  Sea  to  English 
Channel.     In  low  water  to  75.5  fathoms. 

Genus  Petaloproctus  Quatrefages,  1865 

Petaloproctus  tenuis  (Theel,  1879) 

Figure  34,  I,  m 

Maldane  tenuis  Th^el,  1879,  p.  57,  pi.  4,  figs.  52-54. 

Maldane  filifera  Verrill,  1879,  p.  179. 

Petaloproctus  tenuis  Arwidsson,  1907,  p.  114,  pi.  6,  fig.  190a. — Annenkova,  1937, 

p.  181;  1938,  p.  193.— Zatsepin,  1948,  p.  145,  pi.  34,  fig.  5.— Wesenberg-Lund, 

1948,  p.  31.— Berkeley  and  Berkeley,  1952,  p.  55,  fig.  113. 
Petaloproctus  tenuis  var.  borealis  Arwidsson,  1907,  p.  118,  pi.  3,  figs.  85-90;  pi.  8, 

figs.  268-272.— Eliason,  1920,  p.  66.— Hartman,  1948,  p.  42.— Berkeley  and 

Berkeley,  1952,  p.  56,  figs.  114-116. 
Petaloproctus  filifer  Hartman,  1942b,  p.  131,  fig.  11,  a-b. 


MARINE    POLYCHAETE    WORMS — PETTIBONE  307 

Description. — Length  18  mm.,  width  0.5  mm.,  20-21  setigerous 
segments,  smgle  achaetous  antanal  segment  (may  appear  as  two  or 
three  on  longer  ventral  side).  Prostomium  rounded,  with  convex 
median  ridge  and  anteroventraUy  extended  brim.  First  three  setigers 
with  one  or  two  acicular  nem"0setae;  with  some  long,  threadlike,  sinu- 
ous notosetae  covered  with  spinules.  Pygidium  with  obUque,  slightly 
concave,  disclike,  asymmetrical  plate  (absent  dorsally),  wide  on  ven- 
tral side,  with  rim  smooth  or  slightly  crenulate;  anal  opening  terminal. 

Remarks. — The  Point  Barrow  specimen  agrees  with  the  stem  form 
of  P.  tenuis,  with  20  setigerous  segments  and  the  rim  of  the  pygidial 
disc  slightly  crenulate  (in  var.  horealis  and  Jilifer  there  are  21  setigers 
and  the  rim  of  the  pygidial  disc  is  smooth) , 

Nev)  records. — Arctic  Alaska:  Off  Point  Barrow  base,  16  miles 
from  shore,  78.2  fms.,  on  bottom  of  worm  tubes,  few  rocks  (1  station, 
1  specimen).  Canadian  Arctic:  Ducketts  Cove,  Hurd  Channel, 
Melville  Peninsula,  1-14  fms.,  in  sandy  tubes  on  empty  tubes  of 
Pectinaria,  Bartlett,  1933.  Greenland:  Small  bay.  Camp  No.  2, 
Nugsuak  Peninsula,  74°7'  N.,  on  empty  tubes  of  Pectinaria,  J.  C. 
Martin,  1897. 

Distribution. — Scattered  records  in  the  Arctic:  Alaskan  and  Cana- 
dian Arctic,  East  Greenland,  Spitsbergen,  Novaya  Zemlya.  Also  off 
Cape  Cod,  Massachusetts;  southeastern  Alaska  to  British  Columbia; 
north  Japan  Sea.    In  1-833  fathoms. 

Family  Sabellariidae 

Body  divided  into  three  regions:  (1)  anterior  thoracic  region  includ- 
ing two  anterior  thoracic  segments  with  parapodia  reduced  to  neuro- 
podia  with  capillary  setae  and  two  large,  more  or  less  fused,  heavy 
columns,  the  opercular  peduncles,  directed  anteriorly,  bearing  very 
modified  setae  arranged  in  one  to  three  concentric  rows  and  forming  an 
operculum  for  closing  opening  of  tube;  three  or  four  para  thoracic 
biramous  segments  with  capillary  neurosetae  and  paddlelike  or 
styliform  notosetae  and  dorsal  falciform  branchiae  (fig.  34,  n-q) ;  (2) 
abdominal  region  with  notopodia  in  form  of  broadly  flattened  pinnules 
bearing  pectinate  uncini,  and  neuropodia  with  capillary  setae;  with 
dorsal,  simple,  ligulate  or  falciform  branchiae;  (3)  caudal  region, 
achaetous,  apodous,  cylindrical,  without  appearance  of  segmentation. 
Prostomium  indistinct,  between  opercular  peduncles,  with  pair  of 
palps  and  usually  a  median  tentacle.  Mouth  unarmed,  usually  sur- 
rounded with  numerous  filiform  tentacles.  Tube  of  sand  or  fine 
gravel,  concreted,  resistant,  may  be  solitary  or  colonial,  constructing 
sandy  reefs. 

Represented  by  a  single  species  from  Point  Barrow. 


308  PROCEEDINGS    OF   THE   NATIONAL   MUSEUM  voi^  m 

Genus  Idanthyrsus  Kinberg,  1867 
Idanthyrsus  armatus  Eauberg,  1867 

Figure  34,  n-q 

Idanthyrsus  armatus  Kinberg,  1867,  p.  350. — Johansson,  1926,  p.  9;  1927,  p.  90. — 
Monro,  1930,  p.  117,  fig.  73;  1936,  p.  172.— Annenkova,  1937,  p.  184;  1938, 
p.  196.— Okuda,  1938a,  p.  242,  figs.  4,  5.— Hartman,  1944c,  p.  336,  pi.  31, 
fig.  36.— Berkeley  and  Berkeley,  1952,  p.  107,  figs.  220-222. 

Idanthyrsus  ornamentatus  Chamberlin,  1919,  p.  262,  pi.  3,  figs.  2-5. — Hartman, 
1944c,  p.  337,  pi.  31,  fig.  34;  1948,  p.  43.— Hartman  and  Reish,  1950,  p.  41. 

Description. — Length  up  to  60  mm.,  width  5  mm.,  caudal  lobe  11 
mm.  Body  widest  anteriorly,  tapering  gradually  posteriorly,  with 
achaetous  caudal  region  sharply  set  off  from  body.  Opercular  pe- 
duncles elongated,  semicyUndrical,  completely  separated  and  diver- 
gent anteriorly,  obliquely  truncated  distally,  with  two  rows  modified 
golden  setae  or  paleae;  inner  row  paleae  rather  stout,  smooth,  gently 
curved,  tapering  gradually  to  acute  tips,  7-16  per  column;  outer  row 
paleae  more  slender,  straight  or  slightly  curved,  with  coarse  lateral 
spines,  16-36  per  column;  row  of  9-15  distal  papillae  per  column,  at 
base  of  outer  paleae.  One  or  two  pairs  heavy,  deep  amber-colored, 
strongly-bent  nuchal  hooks  on  dorsal  side  opercular  peduncles. 
Numerous  filiform  oral  tentacles  on  ventral  side  opercular  peduncles. 
Pair  of  grooved  slender  palps  and  median  tentacle  between  bases  of 
peduncles.  First  thoracic  segment  short,  with  ventral  group  capiUary 
setae  lateral  to  conical  papilla.  Second  segment  with  lateral  group 
capillary  setae,  three  papillae  and  dorsal  branchiae.  Three  para- 
thoracic  segments  with  rectangular  palletlike  pinnules  with  flattened, 
paddlelike  ilotosetae;  neurosetae  bipinnate  capillary  setae.  Abdom- 
inal segments  with  wide,  flattened  dorsal  pinnules  bearing  pectinate 
uncini;  capillary  barbed  neurosetae.  Caudal  lobe  smooth,  achaetous. 
Paired  dorsal  branchiae  cirriform,  with  transverse  ridges  on  medial 
side,  begin  on  setiger  2,  absent  on  last  few  abdominal  segments. 
Color:  In  life:  Reddish  brown  to  dark  violet  anteriorly,  branchiae 
red,  paleae  golden  yellow.  In  alcohol:  Colorless,  with  purple 
patches  anteriorly.  Tube:  Thick,  of  coarse  sand  grains  neatly  ce- 
mented together,  very  resistant,  one  side  flattened,  attached  to  rocks, 
shells,  crustaceans;  upper  surface  convex. 

Remarks. — /.  ornamentatus  Chamberlin  is  herein  referred  to  /. 
armatus,  following  Okuda  (1938).  According  to  observations  by  G. 
E.  MacGinitie  it  is  a  hardy  species,  a  specimen  living  in  a  pail  in  the 
laboratory  for  three  days,  on  a  rock  covered  with  a  mass  of  barnacles, 
etc.  Male  specimen,  collected  from  70  fathoms,  spawned  in  the 
laboratory  August  9,  1949. 


MARINE    POLYCHAETE    WORMS — PETTIBONE  309 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  up  to  7 
miles  from  shore,  4.3-70  fms.,  on  bottoms  of  mud,  and  gravel,  stones 
(3  stations,  6  specimens) .  Bering  Sea  :  Bering  Island,  N.  Grebiatzky, 
1884.  West  Coast  North  America:  Albatross  Sta.  2842,  54°15'  N., 
166°3'  W.,  72  fms.;  Washington  and  Puget  Sounds,  10-110  fms., 
Pettibone. 

Distribution. — Arctic  Alaska,  Bering  Sea  to  Panamd;  north  Japan 
Sea  to  Japan;  west  coast  South  America  (ChUe),  Straits  of  Magellan, 
Falkland  Islands.     In  low  water  to  110  fathoms. 

Family  Sternaspidae 

Body  very  short,  grublike,  aberrant;  segments  few,  uniramous  or 
achaetous,  without  parapodial  lobes  (fig.  35,  a).  Prostomium  re- 
duced to  small  rounded  tubercle,  without  appendages.  Mouth 
rounded,  subterminal,  ventral  to  prostomium.  Body  divided  into 
three  regions:  (1)  anterior,  with  complete  rings,  with  lateral  concentric 
rows  of  strong  acicular  yeUow  setae  on  anterior  three  segments 
(anterior  region  may  be  retracted  into  more  posterior  segments); 
(2)  middle,  with  segmental  divisions  absent  on  midventral  part, 
without  setae  visible  externally;  (3)  posterior,  with  ventral  horny 
shield  formed  by  two  trapezoidal  plates  provided  on  their  external 
sides  with  radiating  bundles  of  stiff,  barbed  and  smooth  capillary 
setae  (fig.  35,  b).  Numerous,  long,  filiform  anal  branchiae,  inserted 
on  posterior  end.     Anus  terminal.     Lives  in  mud. 

Represented  by  a  single  species  from  Point  Barrow  (the  family  is 
usually  considered  to  have  a  single  cosmopolitan  species). 

Genus  Sternaspis  Otto,  1821 
Stemaspis  scutata  (Ranzani,  1817) 

Figure  35,  a,  b 

Thalassema  scutata  Ranzani,  1817,  p.  1457,  pi.  11,  figs.  10-13. 

Sternaspis  fossor  Stimpson,  1854,  p.  29,  pi.  2,  fig.  19. — Webster  and  Benedict, 
1884,  p.  725.— Moore,  1909b,  p.  144;  1923,  p.  218.— Hartman,  1944a,  pp.  336, 
342,  pi.  33,  fig.  15.— Berkeley  and  Berkeley,  1952,  p.  59,  fig.  123. 

Sternaspis  islandica  Malmgren,  1867,  p.  87,  pi.  14,  fig.  85. 

Sternaspis  scutata  Moore,  1903,  p.  487;  1908,  p.  357;  1923,  p.  218.— Fauvel,  1927, 
p.  216,  fig.  76,  a-g;  1932,  p.  213;  1933,  p.  52;  1934a,  p.  60.— Monro,  1930, 
p.  178.— Annenkova,  1937,  p.  185;  1938,  p.  196.— Hartman,  1942c,  p.  102.— 
Gorbunov,  1946,  p.  39.— Wesenberg-Lund,  1949,  p.  345;  1950a,  p.  46;  1950b, 
p.  104;  1951,  p.  98.— Hartman  and  Reish,  1950,  p.  38. 

Sternaspis  scutata  var.  africana  Monro,  1930,  p.  179. 


310  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  los 

Description. — Up  to  31  mm.  long,  14  mm.  wide.  Body  variable  in 
shape;  may  be  inflated  at  both  ends,  anterior  region  may  be  retracted 
into  more  posterior  segments.  Integument  densely  pilose,  covered 
with  fine  filiform  papillae.  Anterior  region  of  seven  segments; 
segments  1  and  5-7  achaetous;  segments  2-4  each  with  lateral  semi- 
circular row  of  strong,  short,  yellow  setae  which  are  arched,  pointed 
to  blunt  (worn  down),  diminishing  in  size  toward  ventral  side;  pair 
of  long  genital  papillae  on  segment  7.  Middle  region  of  six  segments, 
weU  defined  except  for  narrow  midventral  strip,  with  bundles  of  capil- 
lary setae  embedded  in  body  wall.  Posterior  region  with  five  or  more 
segments,  with  ventral  horny  trapezoidal  plates  showing  concentric 
and  radiating  striae,  each  plate  with  16  radiating  bundles  of  stiff 
setae,  9  on  lateral  and  7  on  posterior  borders.  Anal  filaments  or 
branchiae  very  numerous,  long,  filiform,  sometimes  spiralled.  Colok  : 
In  alcohol:  Without  color  or  brownish,  with  rusty-red  ventral  shield. 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  up  to  5 
miles  from  shore,  1.7-24.7  fms.,  on  bottoms  of  mud  and  gravel  with 
mud  (3  stations,  5  specimens).  Bering  Sea:  63°37'  N.,  165°19'  W., 
12  fms.,  Stoney,  1884;  off  mouth  Yukon  Eiver,  3  fms.,  E.  W.  Nelson, 
1877.  Alaska:  Iluiliuk  Harbor,  Unalaska,  Dall,  1871;  Albatross 
Sta.  3340,  55°26'  N.,  155°26'  W.,  695  fms.,  and  Sta.  3330,  54°  N., 
166°53'  W.,  351  fms.,  1890.  Chiachi  Islands,  20  fms.,  1874;  off 
Round  Island,  Coal  Harbor,  Unga  Island,  6-8  fms.,  1872;  Eastern 
Harbor,  Sitka,  15  fms.,  gravelly-mud,  1874;  all  collected  by  Dall. 
West  Coast  North  America:  Washington  and  Puget  Sounds, 
10-111  fms.,  mud,  Pettibone.  East  Coast  North  America:  Nova 
Scotia,  Maine,  Massachusetts,  5-102  fms.,  U.  S.  Fish  Commission. 

Distribution. — Scattered  records  in  the  Arctic :  Siberian  and  Alaskan 
Arctic,  Greenland.  Also  Iceland  to  English  Channel,  Mediterranean, 
Adriatic,  Gulf  of  Oman,  Persian  Gulf;  Nova  Scotia  to  Massachusetts; 
Bering  Sea  to  Panamd;  north  Japan  Sea  to  Japan,  China.  Southern 
latitudes:  South  Shetlands,  Palmer  Archipelago,  South  Orkneys, 
South  Georgia,  west  Africa,  Indian  Ocean,  Australia,  New  Zealand. 
In  1.7-766  fathoms. 

Family  Pectinariidae  (Amphictenidae) 

Body  short,  conical,  with  segments  few  in  number.  Prostomium 
indistinct,  fused  with  buccal  segment  forming  a  truncate  anterior  end 
provided  with  two  bundles  of  paleae  (}&Tge,  flattened,  golden  setae) 
in  a  horizontal  row  directed  obliquely  anteriorly,  forming  an  operculum 
for  the  tube  or  as  a  fork  for  digging  in  sand  or  mud ;  ventral  to  paleae, 
an  antennular  or  tentacular  membrane  bordered  or  not  with  short  to 
long  filiform  papillae,  surrounding  numerous  prehensile  oral  tentacles 


MARINE    POLYCHAETE    WORMS — PETTIBONE  311 

not  retractile  in  mouth;  dorsal  and  posterior  to  paleae  a  semilunar 
flat  or  concave  cephalic  plate  limited  posteriorly  by  an  entire  or 
crenulate  rim;  two  pairs  tentacular  cirri  with  first  pair  lateral  to  base 
of  paleae,  second  pair  lateral  to  rim  of  cephalic  plate  (fig.  35,  c,  d). 
Body  divided  into  three  regions:  (1)  anterior  thoracic,  including  two 
achaetous  segments  each  with  a  pair  of  lateral  pectinate  branchiae, 
three  notopodial  uniramous  setigers  lacking  neuropodia;  ventral  side 
anterior  region  with  thick  glandular  cushions ;  (2)  median  abdominal, 
with  biramous  segments  (except  last  few  may  have  notosetae  only), 
notopodia  with  setae  capillary,  limbate,  smooth  or  finely  den  tided; 
neuropodia  in  form  of  wide  pinnules  bearing  single  row  of  pectiniform 
uncini;  (3)  posterior  anal  plaque  or  scapha,  small,  foliaceous,  concave 
dorsally,  folded  under  abdomen,  with  rudimentary  achaetous  segments 
except  for  the  first  segment  which  has  a  series  of  acicular,  more  or 
less  recurved  setae  (the  scaphal  hooks);  an  oval  or  filiform  ligule 
above  the  anus  (fig.  35,  g,  h).  Tube  free,  conical,  rigid,  fragile,  open 
at  both  ends,  straight  or  arcuate,  formed  of  a  single  layer  of  cemented 
sand  grains  or  shells,  and  liued  by  a  membrane.  Live  in  sand  or  mud, 
with  anterior  and  larger  end  of  tube  directed  below.  May  secrete 
mucus  in  abundance. 

Represented  by  a  single  genus  (and  subgenus)  and  two  species 
from  Point  Barrow. 

Genus  Pectinaria  Laxnark,  1818 

Subgenus  Cistenides  Malmgren,  1865 

Both  species  have  the  tube  sHghtly  arched,  lined  with  rusty- 
colored  membrane.  Tentacular  membrane  free,  the  lateral  portions 
not  fused  to  paleal  segment,  with  30-45  filiform  to  clavate  marginal 
papillae.  Oral  tentacles  short,  thick,  grooved.  Dorsal  rim  of  flat 
paleal  segment  entire,  not  den  tided  (fig.  35,  d).  Usually  17  segments 
with  capillary  notosetae  (15-17,  the  last  two  segments  may  have  the 
notosetae  few  in  number  or  lacking),  of  which  12  (beginning  on  setiger 
4)  have  elongate  flattened  pinnules  bearing  single  rows  of  neuropodial 
uncini;  pectiniform  uncini  thin,  flat,  with  3-4  major  teeth  and  4-5 
small  ones  above  basal  process  (fig.  35,  /).  Anal  plaque  distinctly 
separated  from  abdomen  by  constriction,  suboval,  festooned  laterally 
(rudimxcntary  parapodia) ;  basally  with  7-1 1  pairs  short,  heavy  scaphal 
hooks  (fig.  35,  g). 

Key  to  the  species  of  Pectinaria  (Cistenides)  from  Point  Barrow 

1.  Cephalic  paleae  usually  7-9  pairs  (7-13),  heavy,  with  blunt  tips  (or  very  short 

sharp  tips;  fig.  35,  i,  f) P.  (C.)  granulata 

Cephalic  paleae  usually  11-14  pairs  (9-15),  long,  tapered  to  slender  pointed 
tips  (fig.  35,  e) P.  (C.)  hyperborea 


312  PROCEEDINGS    OF   THE   NATIONAL   MUSEUM  voL.m 

Pectinaria  iCistenides)  granulata  (Linne,  1767) 

Figure  35,  i-k 

Sabella  granulata  Linn6,  1767,  p.  1268. 

Cistenides  granulata  Malmgren,  1865,  p.  359. — Webster  and  Benedict,  1887,  p. 

747.— Chamberlin,   1920,  p.  25.— Berkeley  and  Berkeley,   1942,  p.  200.— 

Hartman,  1944a,  pp.  335,  342.— Wesenberg-Lund,  1950b,  p.  105;  1951,  p.  99. 
Pectinaria  brevicoma  Johnson,  1901,  p.  423,  pi.  15,  figs.  151-156. — Not  Moore, 

1923,  p.  216  (= P.  calif orniensis  Hartman;  in  USNM). 
Pectinaria  (Cistenides)  granulata  Hessle,  1917,  p.  77. — Nilsson,  1928,  p.  28,  fig. 

8.— Annenkova,  1937,  p.  186;  1938,  p.  198.— Treadwell,  1937,  p.  32. 
Cistenides  brevicoma  Hartman,  1941b,  p.  331,  pi.  50,  figs.  13,  14,  16;  pi.  52,  fig. 

23;  1944b,  p.  268. 
Pectinaria  (Cistenides)  brevicoma  Berkeley  and  Berkeley,  1952,  p.  106,  figs.  218, 

219. 

Description. — Body  20-52  mm,  long,  6-8  mm.  wide.  Tube  38-52 
mm.  long,  7-8  mm.  in  greatest  diameter,  of  rather  coarse  sand  grains, 
nearly  uniform  in  size.  Paleae  usually  8  or  9  pairs  (7-13),  short, 
heavy,  with  tips  blunt  (may  have  short,  sharp  tips).  Scaphal  hooks 
strongly  hooked,  with  a  distract  shoulder.  Color:  In  life:  Colorless, 
transparent,  with  reddish-orange  internal  organs,  red  branchiae, 
golden  paleae. 

New  records. — Arctic  Alaska:  Eluitkak  Pass,  Elson  Lagoon  near 
Point  Barrow,  stony;  Point  Barrow  base,  washed  ashore;  off  Point 
Barrow  base,  up  to  12.1  miles  from  shore,  3.7-123.5  fms.,  on  bottoms 
of  mud,  stones,  mass  of  worm  tubes,  and  various  combinations  of 
mud,  sand,  gravel,  rocks,  stones,  large  perforated  rocks,  shells  (17 
stations,  53  specimens);  10  miles  west  Point  FrankHn,  13.5  fms., 
sand,  Point  Barrow  Expedition,  1883.  Kamchatka:  Petropavlovsk, 
Grebintzky,  1883.  Bering  Sea:  Atka  Island,  Aleutians,  Turner,  1879; 
Bering  Straits,  13  fms.,  DaU,  1880.  Alaska:  Chichagof  Harbor,  Attn 
Island,  5-7  fms.,  1873;  Coal  Harbor,  Unga  Island,  1872;  and  Port 
Etches,  12-18  fms.,  1874;  all  collected  by  Dall.  Kodiak,  Fisher; 
Wrangel,  Jones.  Washington:  Puget  Sound,  39-83  fms.,  mud,  rocks, 
shells,  Pettibone.  Canadian  Arctic:  Kneeland  Bay,  Frobisher 
Bay,  Baffin  Island,  and  off  Daniels  Island,  Newell  Sound,  Frobisher 
Bay,  10-30  fms.,  Bartlett,  1942.  Northwest  Greenland:  Off 
Conical  Eock,  76°  N.,  67°30'  W.,  20-40  fms.,  1938;  between  Parker 
Snow  Bay  and  Conical  Rock,  25-45  fms.,  1940;  west  side  Wolstemholm 
Island,  12  fms.,  1940;  all  collected  by  Bartlett.  East  Coast  North 
America:  Off  Labrador,  6-12  fms.,  Bliie  Dolphin  Expeditions,  1949, 
1950, 1951;  off  New  Brunswick,  Nova  Scotia,  Maine,  New  Hampshire, 
Massachusetts,  Long  Island  Sound,  5-190  fms.,  U.  S.  Fish  Commission. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian,  Alaskan, 
and  Canadian  Arctic,  Greenland,  Spitsbergen,  Novaya  Zemlya.  Also 


MARINE    POLYCHAETE    WORMS — PETTIBONE 


313 


J) 


K 


y 


J 


Figure  35. — Sternaspidae:  a,  Sternaspis  scutata,  ventral  yiew  anterior  end;  h,  same,  ventral 
view  posterior  end.  Pectinariidae:  c,  Pectinaria  hyperhorea,  ventral  view  anterior  end; 
d,  same,  dorsal  view  anterior  end;  e,  same,  palea;/,  same,  pectiniform  uncinus;  g,  same, 
dorsal  view  posterior  anal  plaque  or  scapha;  h,  same,  scaphal  seta;  i,  Pectinaria  granulata, 
cephalic  paleae;  /,  same,  cephalic  paleae,  with  short  mucronate  tips;  ^,  same,  scaphal 
hook.     (For  explanation  of  symbols,  see  p.  210.) 


314  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  103 

Iceland,  Faroes;  Labrador  to  Long  Island  Sound;  Bering  Sea  to  western 
Mexico;  north  Japan  Sea.     In  low  water  to  190  fathoms. 

Pectinaria  (Cistenides)  hyperborea  (Malmgren,  1865) 

FiGUKE  35,  c-h 

Cistenides  hyperborea  Malmgren,  1865,  p.  360,  pi.  18,  fig.  40. — Moore,  1903,  p.  479. — 
Fauvel,  1914,  p.  277,  pi.  26,  figs.  27,  28.— Augener,  1928,  p.  775.— Berkeley 
and  Berkeley,  1942,  p.  201. — Wesenberg-Lund,  1950a,  p.  46;  1950b,  p.  106; 
1951,  p.  100. 

Pectinaria  (Cistenides)  hyperborea  Hessle,  1917,  p.  76. — Nilsson,  1928,  p.  31,  fig. 
9.— Gustafson,  1936,  p.  8.— Okuda,  1937b,  p.  56,  fig.  5;  pi.  2,  fig.  F.— Annen- 
kova,  1937,  p.  186;  1938,  p.  197.— Zatsepin,  1948,  p.  147,  pi.  37,  fig.  4. 

Description. — Body  28-55  nun.  long,  6-12  mm.  wide.  Tube  45-72 
mm.  long,  7-10  mm.  in  greatest  diameter,  formed  of  fine  sand  grains 
plus  some  coarser  ones.  Paleae  usually  11-14  pairs  (9-15),  long, 
tapered  to  slender  tips.  Scaphal  hooks  taper  gradually,  without 
distinct  shoulder. 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  up  to  15 
miles  from  shore,  6-123.5  fms.,  on  bottoms  of  mud,  mass  of  worm 
tubes,  and  various  combinations  of  mud,  gravel,  stones,  rocks,  with 
shells,  worm  tubes;  from  fish  trap  (8  stations,  19  specimens).  Bering 
Sea:  Albatross  Sta.  3610,  55°58'  N.,  167°16'  W.,  75  fms.,  mud,  1895. 
Alaska:  Albatross  Sta.  4244,  Kasaan  Bay,  Prince  of  Wales  Island, 
50-54  fms.  West  Greenland:  Godhavn,  U.S.S.  Alert,  1884.  East 
Coast  North  America:  Off  Labrador,  8-125  fms..  Blue  Dolphin 
Expeditions,  1949,  1950,  1951;  off  Nova  Scotia,  Maine,  Massachusetts, 
16-65  fms.,  U.  S.  Fish  Commission. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian  and 
Alaskan  Arctic,  Greenland,  Spitsbergen,  Novaya  Zemlya,  Kara  Sea. 
Also  Iceland,  Norway  to  Danish  waters.  North  Sea;  Labrador  to 
Massachusetts;  Bering  Sea  to  Alaska;  north  Japan  Sea  to  Japan. 
In  1.5-379  fathoms. 

Family  Ampharetidae 

Prostomium  (or  tentacular  membrane)  more  or  less  distinctly 
trilobed,  with  numerous  filiform  oral  tentacles,  smooth  or  pinnate, 
retractile  in  mouth  (fig.  36,  a,  b,  e).  Branchiae  filiform  or  subulate 
(rarely  pinnate),  two  to  four  pairs,  inserted  on  dorsal  part  of  first 
setigerous  segments.  First  few  segments  achaetous,  with  or  without 
special  group  of  setae  or  paleae  anterior  to  branchiae.  Body  divided 
into  two  distinct  regions:  (1)  thoracic;  conical  notopodia  with  smooth, 
limbate  capillary  setae  and,  beginning  on  setigers  3  or  4,  with  neuro- 
podial  flattened  pinnules  bearing  a  row  of  pectiniform  uncini;  (2) 
abdominal;  uncinigerous  pinnules  only.     With  or  without  dorsal. 


MARINE    POLYCHAETE    WORMS — PETTIBONE  315 

ventral  or  anal  cirri.     Tube  membranous,  covered  with  mud  or  ag- 
glutinated foreign  bodies. 

Eepresented  by  two  genera  and  four  species  at  Point  Barrow. 
Both  genera  have  prostomia  trilobed,  without  projecting  glandular 
crests;  oral  tentacles  numerous,  pinnate;  without  large  dorsal  hooks 
posterior  to  branchiae;  branchiae  four  pairs,  filiform  or  subulate, 
arising  on  or  near  a  distinct  transverse  dorsal  ridge;  thoracic  setigers 
14;  uncinigerous  pinnules  begin  on  setiger  3;  notopodia  without 
claviform  cirri. 

Key  to  the  genera  of  Ampharetidae  from  Point  Barrow 

1.     With  paleae,  lateral  group  of  golden  setae  anterior  to  branchiae   (fig.  36, 

o,  b) Ampharete  (p.  315) 

Without  paleae Asabellides  (p.  318) 

Genus  Ampharete  Malmgren,  1865 

All  three  species  have  prostomium  distinctly  trilobed;  median  lobe 
with  pair  of  basal  eyes,  lateral  lobes  encircling  median  lobe  posteriorly 
(fig.  36,  a).  First  two  segments  achaetous;  first  or  buccal  segment 
extended  anteroventrally  forming  large  lower  lip.  Segment  3  or 
paleal  segment  with  group  of  paleae  an*anged  in  semicircle  just  ante- 
rior and  lateral  to  branchiae.  Abdominal  segments  with  neuropodial 
cirri  dorsally  on  uncinigerous  pinnules  poorly  developed  (short, 
rounded  to  conical)  to  well  developed  (cirrif orm,  longer  than  pinnule) . 

Key  to  the  species  of  Ampharete  from  Point  Barrow 

1.  First  2  abdominal  segments  with  enlarged,   padlike,   achaetous   notopodial 

lobes A.  vega 

Without  enlarged  notopodial  lobes  on  first  few  abdominal  segments 2 

2.  Paleae  slender,  taper  gradually.     Abdominal  setigers  12,  usually  with  promi- 

nent neuropodial  cirri.     Anal  cirri  numerous  (fig.  36,  c) A.  acutifrons 

Paleae  taper  rather  abruptly,  with  short  mucronate  tips.     Abdominal  setigers 
16-17,  with  neuropodial  cirri  inconspicuous.    Anal  cirri  2 A.  goesi 

Ampharete  vega  (Wiren,  1883) 

Figure  36,  a 

Amphideis  vega  Wir6n,  1883,  p.  415,  pi.  32,  figs.  3,  4. 

Ampharete  vega  Hessle,  1917,  p.  99. — Augener,  1928,  p.  778. — Annenkova,  1929, 
p.  493.— Zatsepin,  1948,  p.  150. 

Description. — Length  16  mm.,  width  1.5  mm.  (up  to  50  mm.  long, 
4  mm.  wide — Wiren,  1883).  Paleae  slender,  tapering  gradually  to 
fine  tips,  10-13  in  each  group.  Eight  branchiae  form  continuous 
transverse  group,  arising  from  low  fold.  Abdominal  segments  up  to 
27-28  (16  in  Point  Barrow  specimen;  complete?).     First  two  abdom- 

261112—54 8 


316  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  103 

inal  segments  with  enlarged,  rounded,  padlike,  achaetous  notopodial 
lobes.  Neuropodial  cirri  poorly  developed  or  lacking.  Pygidium  with 
several  small  papillae?  Colorless  in  alcohol.  Tube  thin,  membra- 
nous, with  sand  and  debris,  rather  straggly. 

New  records. — Arctic  Alaska:  West  side  Elson  Lagoon  near  en- 
trance to  small  lagoon  to  west,  near  Point  Barrow  (rather  brackish 
water),  1.2  fms.  (1  station,  1  specimen). 

Distribution. — Scattered  records  in  the  Arctic :  Siberian  and  Alaskan 
Arctic,  Spitsbergen,  Barents  Sea,  Kara  Sea,  Laptev  Sea.  In  L2-11 
fathoms. 

Ampharete  acutifrons  (Grube,  1860) 

Figure  36,  b-d 

Amphideis  acutifrons  Grube,  1860,  p.  109,  pi.  5,  fig.  6. 

Ampharete  grubei  Malmgren,  1865,  p.  363,  pi.  19,  fig.  44. — Eliason,  1920,  p.  70. — 

Fauvel,  1927,  p.  227,  fig.  79,  a-p.— Augener,  1928,  p.  776.— Thorson,  1946, 

p.  121,  fig.  66.— Hartman,  1948,  p.  43. 
Ampharete  cirrata  Webster  and  Benedict,  1887,  p.  747,  pi.  8,  figs.  110-112. 
Ampharete  trilobata  Webster  and  Benedict,  1887,  p.  747. 
Ampharete  acutifrons  Hessle,   1917,  p.  96. — Annenkova,  1937,  p.  188;  1938,  p. 

200.— Zatsepin,  1948,  p.  150,  pi.  37,  fig.  8.— Wesenberg-Lund,  1950a,  p.  47; 

1950b,  p.  109;  1951,  p.  102. 

Description. — Length  45  mm.,  width  6  mm.  (up  to  56  mm.  long, 
8  mm.  wide — Wiren,  1883).  Paleae  slender,  taper  gradually  to  long, 
delicate  tips,  may  be  somewhat  curved,  10-30  in  each  group.  Bran- 
chiae in  two  groups  separated  middorsally,  each  group  with  three 
branchiae  in  transverse  line  and  one  slightly  more  posterior.  Abdom- 
inal segments  12.  With  neuropodial  cirri  small,  conical  to  quite 
long,  longer  than  pinnule  (weU  developed  on  Point  Barrow  specimens). 
Pygidium  a  short  ring  bearing  a  circle  of  numerous  subequal  anal 
cirri  (2  lateral  ones  may  be  longer).  Colorless  in  alcohol.  Tube 
membranous,  covered  with  soft,  gray  debris  (or  agglutinated  mud). 

Remarks. — The  types  of  Ampharete  cirrata  Webster  and  Benedict 
from  Eastport,  Maine,  were  examined.  This  species  was  separated 
on  the  basis  of  the  relative  development  of  the  neuropodial  cirri, 
which  appears  to  be  somewhat  variable.  The  types  of  A.  trilobata 
Webster  and  Benedict,  also  from  Eastport,  were  not  available,  but 
this  species  also,  according  to  the  original  description,  was  differen- 
tiated by  the  relative  development  of  the  cirri. 

New  records. — Arctic  Alaska:  Eluitkak  Pass,  Elson  Lagoon  near 
Point  Barrow;  Point  Barrow  base,  washed  ashore;  off  Point  Barrow 
base,  up  to  6  miles  from  shore,  3.7-49  fms.,  on  bottoms  of  mud,  and 
various  combinations  of  mud,  pebbles,  gravel,  stones,  rocks,  and 


MARINE    POLYCHAETE    WORMS — PETTIBONE  317 

shells  (10  stations,  17  specimens).  West  Coast  North  America: 
Washington  and  Puget  Sounds,  80  fms.,  Pettibone.  East  Coast 
North  America:  Off  Labrador,  8  fms,,  Blue  Dolphin  Expedition, 
1949. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian  and 
Alaskan  Arctic,  Greenland,  Jan  Mayen,  Spitsbergen,  Novaya  Zem- 
lya.  Also  Iceland,  Swedish  west  coast  to  France,  Mediterranean; 
Labrador  to  Maine;  Bering  Sea  to  southern  California;  north  Japan 
Sea.     In  1-1,333  fathoms. 

Ampharete  goesi  Malmgren,  1865 

Ampharete  goesi  Malmgren,  1865,  p.  364,  pi.  19,  fig.  45. — Hessle,  1917,  p.  91. — 
Augener,  1928,  p.  778.— Annenkova,  1929,  p.  492,  pi,  38,  fig,  37;  1937,  p.  188; 
1938,  p.  200.— Zatsepin,  1948,  p.  150,  pi.  38,  fig.  10.— Wesenberg-Lund, 
1950a,  p.  47;  1950b,  p.  110;  1951,  p.  102.— Berkeley  and  Berkeley,  1952,  p. 
66,  fig.  136. 

Ampharete  goesi  subsp.  braznikovi  Annenkova,  1929,  p.  492,  pi.  38,  fig.  44. 

Description. — Length  35  mm,,  width  6  mm.  (up  to  50  mm.  long,  7 
mm.  wide — Malmgren,  1865).  Paleae  rather  stout,  taper  abruptly, 
ending  in  short  acuminate  tips,  14-21  in  each  group.  Branchiae  in 
two  groups  separated  middorsally,  each  group  with  three  branchiae 
in  transverse  line.  Abdominal  segments  17  (stem  form  and  Point 
Barrow  specimens)  or  16  (subsp.  braznikovi  and  Washington  Sound 
specimens).  Neuropodial  cirri  inconspicuous,  low,  rounded.  Pygid- 
ium  with  two  long  anal  cirri.  Color:  In  life:  Body  reddish  orange, 
branchiae  green.  In  alcohol:  Colorless.  Tube:  Membranous,  rather 
thick,  with  debris  including  bits  of  shells,  sea  urchins  tests,  and 
foraminiferans,  very  straggly. 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  up  to  15 
miles  from  shore,  20.3-123.5  fms.,  on  bottoms  of  mass  of  worm  tubes 
and  various  combinations  of  mud,  pebbles,  gravel,  stones,  rocks,  shells, 
and  worm  tubes  (11  stations,  20  specimens).  West  Coast  North 
America:  Washington  Sound,  Pettibone. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian  and 
Alaskan  Arctic,  Greenland,  Spitsbergen,  Barents  Sea,  White  Sea, 
Novaya  Zemlya.  Also  Iceland;  British  Columbia  to  Washington; 
Okhotsk  Sea  to  north  Japan  Sea.     In  10.5-353  fathoms. 

Ampharete  sp.  (Young) 

Two  very  small  specimens,  12  mm.  long,  were  found  on  bottom  of 
rocks  with  bryozoans  and  hydroids  in  17  fathoms  September  9,  1948. 
Prostomium  3-lobed,  with  four  eye  spots,  bases  of  four  pairs  branchiae, 
14  thoracic  segments,  10  abdominal  segments  plus  a  growing  zone. 


318  PROCEEDINGS    OF   THE    NATIONAL  MUSEUM  vol.  103 

GerkusAsabellides  Annenkova,  1929 

Includes  Pseudosahellides  Berkeley  and  Berkeley,  1943. 

Asabellides  sibirica  (Wiren,  1883) 

Figure  36,  e 

Sabellides  sibirica  Wir6n,  1883,  p.  418. 

Asabellides  orientalis  Annenkova,  1929,  p.  494,  pi.  38,  figs.  50,  51,  pi.  39,  figs. 

60-65;  1937,  p.  188. 
Asabellides  sibirica  Annenkova,  1938,  p.  201. 
Pseudosabellides  lineata  Berkeley  and  Berkeley,  1943,  p.  131;  1944,  p.  3;  1952, 

p.  71,  fig.  147. 

Description. — Body  long,  slender,  38  mm.  long,  1.5  mm.  wide. 
Prostomium  trilobed  with  median  lobe  wider  and  flared  anteriorly, 
lateral  lobes  diagonal.  Segments  1  and  2  fused  laterally  and  ventrally, 
forming  large  lower  lip;  dorsally,  appear  as  two  short  segments  more 
or  less  hidden  by  dorsal  branchial  fold.  Segments  3  and  4  more  or  less 
fused  ventrally  and  laterally;  dorsally,  segment  3  without  paleae  but 
with  branchiae;  segment  4  with  first  notopodial  lobes  and,  middorsally, 
a  fused  nephridial  area  wath  pair  of  papillae  posterior  to  branchial 
bases.  Branchiae  slender,  fihform,  emerge  from  prominent  ridge,  six 
in  transverse  row,  two  slightly  posterior.  Abdominal  segments  23 
(14-23).  Neuropodial  cirri  smaU,  short,  rounded  (Point  Barrow 
specimens)  to  long,  slender,  cirriform.  Pygidium  with  few  small  cirri 
and  pair  of  long  lateral  ones.  Color:  In  life:  Salmon  below,  darker 
above,  branchiae  green,  oral  tentacles  pink.    In  alcohol:  Colorless. 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  5  mUes  from 
shore,  11.7  fms.,  on  bottom  of  mud  (1  station,  3  specimens). 

Distribution. — Scattered  records  in  the  Arctic:  Siberian,  Alaskan, 
and  Canadian  Arctic.  Also  Hudson  Bay;  Bering  Sea  to  British 
Columbia;  Okhotsk  Sea  to  north  Japan  Sea.    In  4-106  fathoms. 

Family  Terebellidae 

Body  divided  into  two  regions:  (1)  thoracic;  more  or  less  inflated, 
with  dorsal  bimdles  of  capillary  setae  and  ventral  uncinigerous  tori 
(sometimes  acicular  crotchets,  sometimes  absent),  often  with  ventral 
glandular  shields;  (2)  abdominal;  elongated,  more  or  less  tapering, 
most  often  lacking  dorsal  setae  and  bearing  only  uncinigerous  pinnules. 
Prostomium  of  variable  form  and  dimensions,  fused  with  buccal  seg- 
ment, forming  a  lower  lip,  a  transverse  dorsal  cephalic  ridge  with  or 
without  numerous  eye-spots,  a  tentacular  membrane  with  numerous 
filiform  grooved  tentacles  not  retractile  in  mouth,  a  semicircular  or 
folded  upper  lip  (fig.  36,/).  Branchiae  1-3  pairs,  filiform  or  arbores- 
cent,  or  single,   quadrilobed,   pectinate,   on  anterior  segments,   or 


MARINE    POLYCHAETE    WORMS — PETTIBONE 


319 


Figure  36. — A.mph.arttid&e: a,  Ampharetevega,  dorsal  view  anterior  end;  b,  Ampharete  acuti- 
frons,  dorsal  view  anterior  end;  c,  same,  lateral  view  posterior  end;  d,  same,  palea;  i, 
Asabellides  sibirica,  dorsal  view  anterior  end.  Terebellidae:  /,  lateral  view  anterior  end 
of  terebellid;  g,  Amphitriu  cirraia,  lateral  view  anterior  end;  h,  same,  notoseta;  i,  Nicolea 
venustula,  lateral  view  anterior  end  of  male;  ;,  same,  thoracic  uncini;  k,  Pista  maculata, 
lateral  view  anterior  end,  tentacles  missing;  /,  same,  uncinus  with  long  manubrium.  (For 
explanation  of  symbols,  see  p.  210.) 


320  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  tol.  103 

branchiae  lacking.     Without  paleae.     Tube  usually  membranous, 
encrusted  with  sand,  debris  of  shells,  algae,  etc. 

Represented  by  10  genera  and  11  species  at  Point  Barrow. 

Key  to  the  genera  of  TerebeUidae  from  Point  Barrow 

1.  Uncini  in  single  rows  on  first  6  uncinigerous  segments,  then  in  2  rows  (alter- 

nating, intercogging  or  opposite)  on  certain  number  of  following  segments. 
With  distinct  thoracic  ventral  shields  (single,  not  paired) .  Notosetae  begin 
on  segment  4;  uncini  begin  on  segments  5  or  6  (setigers  2  or  3),  Fused 
prostomium  and  buccal  segment  forming  a  prominent  rectangular  lower  lip, 
a  transverse  semicircular  cephalic  ridge  (with  or  without  numerous  eye-spots 
along  basal  groove),  flared  semicircular  upper  lip,  semicircular  depressed 
tentacular  region  between  upper  lip  and  cephalic  ridge  with  numerous, 

filiform,  grooved  tentacles  (fig.  36,/) 2 

Uncini  in  single  rows  only  (or  with  crotchets  on  thoracic  segments) 7 

2.  With  1-3  pairs  branchiae  dorsally  on  segments  2-4  (fig.  36,  g,  i,  k).     Uncini 

begin  on  segment  5  (setiger  2) 3 

Without  branchiae.     With  lateral  lobes  on  anterior  segments  (fig.  37,  o) 5 

3.  Notosetae   with  tips  finely   denticled    (fig.   36,   h).     Cephalic  ridge  usually 

without  eye-spots.     With  lateral  lobes  on  anterior  segments  (fig.  36,  g). 

Amphitrite  (p.  321) 
Notosetae  with  tips  smooth.     Cephalic  ridge  usually  with  eye-spots 4 

4.  Without  lateral  lobes  on  anterior  segments.     Uncini  of  anterior  segments 

without  long  manubrium  (fig.  36,  j).  Branchiae  2  pairs,  branched  dichoto- 
mously  (fig.   36,  t) Nicolea  (p.  322) 

With  large  lateral  lobes  on  buccal  segment,  enclosing  prostomium  and  con- 
nected ventrally  (fig.  36,  k).  Uncini  of  first  few  segments  with  long  manu- 
brium (fig.  36,  I).     Branchiae  1-3  pairs,  arborescent,  with  main  trunk  well 

marked Pista    (p.    323) 

6.  Uncini  begin  on  segment  6  (setiger  3).  Notosetae  of  2  kinds,  smooth  and 
denticled Proclea  (p.  325) 

Uncini  begin  on  segment  5  (setiger  2).     Notosetae  with  tips  smooth 6 

6.  Uncini  in  single  rows  on  first  6  uncinigerous  segments,  in  double  rows  on  next 

10.     Nephridial  papillae  of  segment  3  not  elongated.     Notosetae  broadly 

bilimbate  below  whipUke  tip  (fig.  37,  b) Leaena  (p.  325) 

Uncini  in  single  rows  on  first  6  uncinigerous  segments,  in  double  rows  on  next 
8.  Nephridial  papillae  of  segment  3  (anterior  to  first  setiger)  elongated. 
Notosetae  weakly  limbate  on  one  side  along  most  of  its  length  (fig.  37,  c). 

Lanassa  (p.  326) 

7.  Uncini  all  short,  avicular  (fig.  37,  d) 8 

Uncini  of  2  types;  thoracic  aciculiform,  with  long  manubrium  (fig.  37,  h); 

abdominal  avicular  (fig.  37,  i).     Without  ventral  shields 9 

8.  Branchiae   2-3   pairs,    cirriform,   in   transverse   rows.     Notosetae   begin   on 

segment  3,  continuing  on  large  number  of  segments;  uncini  begin  on  segment 
5  (setiger  3).     Ventral  shields  usually  slightly  distinct.     Tentacular  lobe 

not  especially  enlarged Thelepus  (p.  327) 

Without  branchiae.  Notosetae  begin  on  segments  2  or  3;  uncini  begin  on 
variable  number  of  segments,  may  be  absent  from  thoracic  region.  With 
ventral  shields,  first  unpaired,  rest  paired,  more  or  less  separated  in  median 
line  (fig.  37,  /).  Tentacular  membrane  large,  trilobed  or  undulating, 
bearing  very  numerous  tentacles  (fig.  37,  e-/), Polycirrus  (p.  328) 


MARINE   POLYCHAETE   WORMS — PETTIBONE  321 

Branchiae  2-3  pairs,  single,  filiform  (fig.  37,  g).  Notosetae  and  uncini 
(crotchets)  both  begin  on  segment  6.  Tentacular  lobe  folded,  with  very- 
numerous  thin,  cylindrical,  and  short,  grooved  tentacles, 

Trichobranchus  (p.  329) 

Branchiae  single,  consisting  of  large  cylindrical  trunk  and  4  pectinate  lobes 
(fig.  37,  j).  Notosetae  begin  on  segment  3,  uncini  begin  on  segment  8 
(setiger  6).  Tentacular  lobe  large,  folded,  with  numerous  tentacles  of  a 
single  kind Terebellides   (p.  330) 


Genus  Amphitrite  O.  F.  Muller,  1771 

Both  species  have  the  body  inflated  anteriorly,  decidedly  atten- 
uated posteriorly.  Cephalic  ridge  without  eye-spots.  Branchiae 
three  pairs,  on  segments  2-4.  Notosetae  begin  on  segment  4;  uncini 
begin  on  segment  5,  in  single  rows  on  first  six,  in  double  rows  on  rest 
of  thoracic  setigers,  in  single  rows  on  abdominal  segments  on  project- 
ing pinnules.     Pygidium  a  crenulate  ring. 

Key  to  the  species  of  Amphitrite  from  Point  Barrow 

1.  Branchiae  cirriform,  a  tuft  of  simple   filaments  from  short,  sUghtly  swollen 

common  base  (fig.  36,  g).     Thoracic  setigers  17 A.  cirrata 

Branchiae  branched.     Thoracic  setigers  19 A.  groenlandica 

Amphitrite  cirrata  Muller,  1776 

Figure  36,  g,  h 

Amphitrite  cirrata  O.  F.  Mtiller,  1776,  p.  216.— Malmgren,  1865,  p.  375,  pi.  21, 
fig.  53.— Webster  and  Benedict,  1887,  p.  748.— Hessle,  1917,  p.  185.— 
Chamberhn,  1920,  p.  22.— Eliason,  1920,  p.  74.— Fauvel,  1927,  p.  251, 
fig.  86,  i-o.— Augener,  1928,  p.  785.— Okuda,  1938b,  p.  102.— Hartman, 
1944a,  pp.  334,  342;  1948,  p.  43.— Thorson,  1946,  p.  127,  figs.  71,  72.— 
Gorbunov,  1946,  p.  39.— Zatsepin,  1948,  p.  156,  pi.  38,  fig.  12.— Wesenberg- 
Lund,  1950a,  p.  50;  1950b,  p.  116;  1951,  p.  106.— Berkeley  and  Berkeley, 
1952,  p.  86,  fig.  175. 

Amphitrite  radiata  Moore,  1908,  p.  350;  1923,  p.  193. 

Description. — Length  up  to  200  mm.,  width  10  mm.  About  12 
ventral  shields.  Branchiae  3  pairs,  cirriform,  a  tuft  of  simple  fila- 
ments from  a  short  common  base.  Each  of  thi-ee  branchial  segments 
(segments  2-4)  with  pair  of  lateral  lobes.  Nephridial  papillae  7 
pairs;  on  segment  3,  prominent  papillae  ventral  to  second  branchial 
pair;  on  segments  6-11,  rather  inconspicuous  papillae  between  noto- 
podia  and  neuropodia.  Thoracic  setigers  17.  Color:  In  alcohol: 
Keddish  brown.  Tube:  Mostly  of  mud,  with  scattered  small  pebbles 
and  debris,  soft,  breaking  easily. 

New  records. — Arctic  Alaska:  Eluitkak  Pass,  Elson  Lagoon;  off 
Point  Barrow  base,  up  to  7  miles  from  shore,  18.3-70  fms.,  on  various 
combinations  of  mud,  sand,  gravel,  stones,  rocks,  large  perforated 
rocks,    shells    (8    stations,    14    specimens).     West    Coast    North 


322  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  103 

America:  Wrangel,  Alaska,  Jones;  Puget  Sound,  low  tide,  Pettibone. 
West  Greenland:  Vaigat,  Disko  Island,  Bartlett,  1937.  East 
Coast  North  America:]  Bay  of  Fundy,  Maine,  Massachusetts, 
14-llO^fms.,  U.  S.  Fish  Commission. 

Distribution. — Widely  distributed  in  the  Arctic :  Siberian,  Alaskan, 
and  Canadian  Arctic,  Greenland,  Spitzbergen,  Franz  Josef  Land, 
White  Sea,  Novaya  Zemlya,  Kara  Sea.  Also  Iceland,  Faroes,  Nor- 
way to  British  Isles,  Azores,  Mediterranean,  Adriatic;  Bay  of  Fundy 
to  Massachusetts;  Bering  Sea  to  southern  California;  Japan.  In  low 
water  to  1,528  fathoms. 

Amphitrite  groenlandica  Malmgren,  1865 

Amphitrite  groenlandica  Malmgren,  1865,  p.  376,  pi.  21,  fig.  52. — Verrill,  1881, 
pp.  305,  310.— Hessle,  1917,  p.  181.— Fauvel,  1927,  p.  250,  fig.  86,  a-c— 
Augener,  1928,  p.  787.— Amaenkova,  1937,  p.  192;  1938,  p.  206.— Gorbunov, 
1946,  p.  39.— Zatsepin,  1948,  p.  157,  pi.  38,  fig.  14.— Wesenberg-Lund, 
1950a,  p.  51;  1950b,  p.  117;  1951,  p.  107. 

Neoamphitrite  robusta  Hartman,  1948,  p.  44  (not  Amphitrite  robusta J ohnaon, 
1901). 

Description. — ^Length  up  to  190  mm.,  width  9  mm.  (a  single,  much 
smaller  specimen  from  Point  Barrow).  About  14  ventral  shields. 
Branchiae  3  pairs,  branched  dichotomously  3-4  times,  with  main 
trunk  short,  thick.  Lateral  lobes  of  first  two  branchial  segments  well 
developed,  continuing  ventrally;  small  lateral  lobes  on  third  branchial 
segment.  Nephridial  papillae  12  pairs  on  segments  3-14,  the  first 
four  with  more  prominent  papUlae.  Thoracic  setigers  19.  Colorless 
in  alcohol.     Tube  of  mud. 

Remarks. — A.  robusta  Johnson  is  very  close  to  A.  groenlandica; 
it  differs  in  having  17  thoracic  setigers  instead  of  19. 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  5  miles  from 
shore,  49  fms.,  on  bottom  of  rocks,  stones,  gravel  (1  station,  1  speci- 
men). Southwestern  Alaska:  Round  Island,  Coal  Harbor,  Unga 
Island,  8-9  fms.,  Dall,  1872. 

Distribution. — Scattered  records  in  the  Arctic:  Siberian  and  Alaskan 
Arctic,  West  Greenland,  Spitsbergen,  Novaya  Zemlya.  Also  Iceland, 
Scandinavian  coast  to  North  Sea,  Ireland;  Maine;  southwestern 
Alaska;  Okhotsk  Sea  to  north  Japan  Sea.     In  7  to  440  fathoms. 

Genus  Nicolea  Malmgren,  1865 
Nicolea  venustula  (Montagu,  1818) 

Figure  36,  i,  j 

Terebella  venustula  Montagu,  1818,  p.  344,  pi.  13,  fig.  2. 

Nicolea  arctica  Malmgren,  1865,  p.  381,  pi.  24,  figs.  66,  67. 

Nicolea  zostericola  Malmgren,  1865,  p.  381,  pi.  26,  fig.  76. — Webster  and  Benedict, 
1887,  p.  749.— Moore,  1909b,  p.  141.— Fauvel,  1927,  p.  261,  fig.  90,  g-n.— 
Annenkova,  1934,  p.  322;  1937,  p.  191;  1938,  p.  205.— Gustafson,  1936,  p. 


MARINE    POLYCHAETE    WORMS — PETTIBONE  323 

9.— Berkeley  and  Berkeley,  1943,  p.  130;  1952,  p.  87,  figs.  177,  178.— Gor- 

bunov,  1946,  p.  39.— Hartman,  1948,  p.  44. 
Nicolea  simplex  Verrill,  1873,  p.  613;  1881,  p.  302,  pi.  10,  fig.  1.— Sumner,  Osburn, 

and  Cole,  1913,  p.  627. 
Nicolea  venustula  Ehlers,  1913,  p.  559. — Hessle,  1917,  p.  171. — Chamberlin,  1920, 

p.  22.— Eliason,  1920,  p.  73.— Fauvel,  1927,  p.  260,  fig.  90,  a-f ;  1934a,  p.  68.— 

Augener,  1928,  p.  788.— Thorson,  1946,  p.  126.— Zatsepin,  1948,  p.  156,  pi. 

38,  fig.  11.— Wesenberg-Lund,  1950a,  p.  51;  1950b,  p.  118;  1951,  p.  109. 

Description. — Length  15-48  mm.,  width  2-5  mm.,  segments  30-46 
(up  to  70  mm.  long — Wiren,  1883).  Ventral  shields  about  14. 
Cephalic  ridge  with  numerous  dark  eye-spots.  Branchiae  two  pairs, 
on  segments  2-3,  branched  dichotomously  3-6  times,  with  very  short 
main  stem,  first  pair  much  larger  than  second.  Without  lateral  lobes 
on  branchial  segments.  Nephridial  papillae  on  segments  3  (slightly 
posterior  to  pair  of  second  branchiae),  6,  and  7;  small  in  female;  in 
male,  papillae  on  segments  6  and  7  long,  cylindrical.  Thoracic  seti- 
gers  15  (14-18,  according  to  Hessle).  Uncini  begin  on  segment  5 
(setigerous  segment  2),  in  single  rows  on  first  six,  in  two  alternating 
rows  on  rest  of  thoracic  segments,  in  single  rows  on  prominent  pro- 
jecting abdominal  pinnules.  Pygidium  crenulate.  Colorless  or 
slightly  brownish  in  alcohol.  Females  with  large  yolky  eggs  inside 
body  (Point  Barrow,  August  21,  and  September  15,  1948;  August  8, 
1949).  Tube  with  tough,  translucent  membranous  lining,  with  small 
pebbles  of  various  sizes,  shell  debris,  foraminiferans,  etc. 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  up  to  12.1 
miles  from  shore,  13.3-123.5  fms.,  on  bottoms  of  mud,  stones,  mass  of 
worm  tubes,  and  various  combinations  of  mud,  sand,  gravel,  pebbles, 
stones,  rocks,  large  perforated  rocks,  with  bryozoans,  hydroids,  shells 
(24  stations,  79  specimens).  Alaska:  Security  Bay,  Jones.  Cana- 
dian Arctic:  Dobbin  Bay,  E.  Ellesmere  Island,  79°36'  N.,  73°35' 
W.,  17  fms.,  Littlewood,  1950.  East  Coast  North  America:  Off 
Maine,  Massachusetts,  Rhode  Island,  Connecticut,  shore  to  63 
fathoms  and  surface  (young),  U.  S.  Fish  Commission. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian,  Alaskan, 
and  Canadian  Arctic,  Greenland,  Jan  Mayen,  Spitsbergen,  Franz 
Josef  Land,  Novaya  Zemlya,  Kara  Sea.  Also  Iceland,  Faroes,  Nor- 
way to  France,  Mediterranean,  Adriatic,  Red  Sea;  Hudson  Bay  to 
Connecticut;  Bering  Sea  to  western  Canada;  north  Japan  Sea;  South 
Africa.     In  low  water  to  472  fathoms. 

Grenus  Pista  Malmgren,  1865 
Pista  maculata  (Dalyell,  1853) 

Figure  36,  k,  I 
Terebella  maculata  Dalyell,  1853,  p.  203,  pi.  28,  figs.  10-14,  19. 
Scione  lobata  Malmgren,  1865,  p.  383,  pi.  23,  fig.  62.— Verrill,  1881,  pp.  305,  310.— 

Webster  and  Benedict,   1887,  p.  749. — Augener,   1928,  p.   789. — Friedrich, 

1939,  p.  127. 


324  PROCEEDINGS   OF   THE    NATIONAL   MUSEUM  vol.  103 

Pista  maculata  Hessle,  1917,  p.  161,  pi.  3,  fig.  4. — Fauvel,  1927,  p.  263,  fig.  91, 
a-h. — Berkeley  and  Berkeley,  1943,  p.  130, — Gorbunov,  1946,  p.  39. — 
Zatsepin,  1948,  p.  155,  pi.  38,  fig.  9.— Wesenberg-Lund,  1950a,  p.  52;  1950b, 
p.  119;  1951,  p.  110. 

Pista  groenlandica  Treadwell,  1937,  p.  33,  figs.  14-16. 

Description. — ^Length  up  to  150  mm.,  width  6  mm.  Ventral  shields 
about  14.  Cephalic  ridge  with  numerous  eye-spots.  Single  pair 
large  branchiae  on  segment  2,  with  large,  cylindrical  main  trunk, 
branched.  Buccal  segment  with  two  large,  rounded  lateral  lobes 
embracing  prostomium,  connected  ventrally  by  a  crest;  segment  2 
short,  without  lateral  lobes;  segment  3  with  large,  rounded,  flattened 
lateral  lobes.  Nephridial  papillae  on  segments  6-7.  Thoracic 
setigers  16  (one  had  small  lobe  on  one  side  only  on  17).  Uncini 
begin  on  segment  5  (setiger  2),  in  single  rows  on  first  6,  in  two  alternat- 
ing intercogging  rows  on  rest  of  thoracic  segments,  in  single  rows  on 
rectangular  pinnules  in  abdominal  region.  Pygidium  with  six  or 
seven  (6-12)  conical  papillae,  arranged  starlike.  Color:  In  alcohol: 
Branchiae  and  tentacles  dark  brown  or  colorless.  Tubes:  Irregularly 
twisted,  with  thin  transparent  membranous  lining  covered  mostly 
with  fine  sand  grains  plus  few  large  pebbles,  bits  of  shell,  bryozoans, 
foraminiferans,  algae,  parts  of  other  worm  tubes. 

Remarks. — The  type  of  P.  groenlandica  Treadwell  from  Baflin  Bay 
was  examined  and  is  herein  referred  to  P.  maculata.  Contrary  to  the 
original  description,  eye-spots  are  present  on  the  cephalic  ridge  (small, 
in  transverse  groove,  easily  overlooked) . 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  up  to  15 
miles  from  shore,  21-123.5  fms.,  on  bottoms  of  mass  of  worm  tubes 
and  various  combinations  of  mud,  gravel,  stones,  rocks,  large  per- 
forated rocks;  from  crab,  Hyas  coarctatus  (13  stations,  168  plus  speci- 
mens). North  Greenland:  5  miles  south  Cape  Chalon;  north 
Omenolu  near  North  Star  Bay,  17  fms.,  Bartlett,  1932.  West 
Greenland:  Oelrichs  Bay,  1937;  off  Conical  Kock,  76°3'  N.,  67°30' 
W.,  20-40  fms.,  1938;  1  mile  northwest  Conical  Rock,  25-60  fms., 
1940;  between  north  shores  Parkers  Snow  Bay  and  Conical  Rock, 
25-45  fms.,  1940;  aU  collected  by  Bartlett.  East  Greenland:  Off 
Cape  Hold  with  Hope,  23^0  fms.,  Bartlett,  1939.  East  Coast 
North  America  :  Off  Labrador,  8-125  fms..  Blue  Dolphin  Expeditions, 
1949,  1950,  1951. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian,  Alaskan, 
and  Canadian  Arctic,  Baffin  Bay,  Greenland,  Spitsbergen,  Barent 
Sea,  Novaya  Zemlya.  Also  Iceland,  Norway  to  English  Channel: 
Hudson  Bay  to  Maine;  Bering  Sea.     In  3-1,528  fathoms. 


AIARINE    POLYCHAETE    WORMS — PETTIBONB  325 

Genus  Proclea  Saint- Joseph,  1894 
Proclea  graffii  (Langerhans,  1884) 

Leaena  graffii  Langerhans,  1884,  p.  262,  pi.  15,  fig.  21. 

Proclea  graffi  Southern,  1914,  p.  120.— Hessle,  1917,  p.  199,  fig.  53.—  Fauvel, 

1927,  p.  268,  fig.  94,  a-g.— Gorbunov,  1946,  p.  39.— Zatsepin,  1948,  p.  157,  pi. 

38,  fig.  19.— Wesenberg-Lund,  1951,  p.  111. 

Description. — Length  up  to  42  mm.,  width  3  mm.,  segments  49. 
Ventral  shields  about  10.  Cephalic  ridge  without  eye-spots.  With- 
out branchiae.  With  lateral  lobes  on  segments  2-4,  those  on  segments 
2  and  3  elongate,  extending  to  ventral  shields.  Nephridial  papillae 
on  segments  3,  6-8.  Thoracic  setigers  16.  Uncini  begin  on  seg- 
ment 6  (setiger  3),  in  single  rows  on  first  six,  in  double  rows  on  rest 
of  thoracic  segments,  in  single  rows  on  projecting  abdominal  pinnules. 
Pygidium  crenulate.     Colorless  in  alcohol.     Tube  ? 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  up  to  7.5 
miles  from  shore,  21-36  fms.,  on  bottoms  of  rocks,  stones,  large  per- 
forated rocks,  from  breaking  rocks  apart  (3  stations,  6  specimens) . 

Distribution. — Scattered  records  in  the  Arctic :  Siberian  and  Alaskan 
Arctic,  Franz  Josef  Land,  White  Sea.  Also  Bering  Sea  to  Okhotsk 
Sea;  Iceland,  Swedish  west  coast,  Finland,  Ireland,  Madeira.  In 
1-36  fathoms. 

Genus  Leaena  Malmgren,  1865 

Leaena  abranchiata  Malmgren,  1865 

Figure  37,  a,  b 

Leaena  abranchiata  Malmgren,  1865,  p.  385,  pi.  24,  fig.  64. — not  Moore,  1909b, 
p.  141  (^Lanassa  venusta). — Ehlers,  1913,  p.  563. — Hessle,  1917,  p.  197, — 
Augener,  1928,  p.  793.— Annenkova,  1937,  p.  192;  1938,  p.  208.— Berkeley 
and  Berkeley,  1943,  p.  130.— Gorbunov,  1946,  p.  39.— Zatsepin,  1948,  p.  157, 
pi.  38,  fig.  17.— Hartman,  1948,  p.  45. —Wesenberg-Lund,  1950a,  p.  53; 
1950b,  p.  121;  1951,  p.  112. 

Leaena  antarctica  Mcintosh,  1885,  p.  462,  pi.  48,  figs.  9,  10;  pi.  28A,  figs.  10,  11. 

Leaena  abranchiata  var.  antarctica  Hessle,  1917,  p.  197. — Monro,  1930,  p.  188; 
1936,  p.  178. 

Description. — Length  43  mm.,  width  3  mm.  (up  to  75  mm.  long,  6 
mm.  wide — Malmgren,  1865).  About  10  ventral  shields.  Cephalic 
ridge  without  eye-spots.  Without  branchiae.  Lateral  lobes  on  seg- 
ment 2  extend  ventrally,  those  on  segment  3  extend  dorsally,  forming 
a  prominent  ridge  (segment  preceding  first  setiger;  ridge  not  so  prom- 
inent in  var.  antarctica).  Thoracic  setigers  10.  Uncini  begin  on  seg- 
ment 5  (setiger  2),  in  single  rows  on  first  6,  in  double  rows  on  next  10, 
then  in  single  rows  on  prominent  abdominal  pinnules.  Notosetae 
widely  bilimbate  below  whiplike  tips.  Colorless  in  alcohol.  Tube  : 
Of  soft  mud  with  a  few  bits  of  scattered  rock  (some  small  ones  on 
tubes  of  Pista  maculata) . 


326  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  103 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  25  fms.,  on 
bottoms  of  gravel,  stones,  shells  (1  station,  1  specimen).  East  Coast 
North  America:  Off  Labrador,  15-55  fms.,  mud  and  mud  with  rock, 
Blue  Dolphin  Expeditions,  1950,  1951. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian  and  Alas- 
kan Arctic,  Baffin  Bay,  Greenland,  Spitsbergen,  Novaya  Zemlya, 
Kara  Sea.  Also  Iceland,  Faroes,  Norway,  Sweden,  Finland;  Hudson 
Bay  to  Labrador;  southwestern  Alaska;  Okhotsk  Sea  to  north  Japan 
Sea.  In  6-833  fathoms.  Var.  antarctica:  Antarctic,  South  Georgia; 
up  to  1,975  fathoms. 

Genus  Lanassa  M almgren,  1865 
Lanassa  venustM  (Malm,  1874) 

Figure  37,  c 

Laphaniella  venusta  Malm,  1874,  p.  98,  pi.  1,  fig.  8. 

Leaena  nuda  Moore,  1905b,  p.  855,  pi.  44,  figs.  14,  15. 

Leaena  abranchiata  Moore,  1909b,  p.  141  (not  L.  abranchiata  Malmgren,  1865). 

Lanassa  venusta  Hessle,  1917,  p.  205. — Eliason,  1920,  p.  75. — Gustafson,  1936, 

p.  9.— Annenkova,  1937,  p.  194;  1938,  p.  209.— Zatsepin,  1948,  p.  157. 
Lanassa  venusta  subsp.  pacifica  Annenkova,  1938,  pp.  209,  230. 

Description. — Length  37-55  mm.,  width  3  mm.,  segments  54-67. 
Ventral  shields  about  10.  Cephalic  ridge  without  eye-spots  (present  in 
subsp.  pacifica).  Oral  tentacles  numerous  (more  than  12  as  indicated 
by  Malm) .  Without  branchiae.  Lateral  lobes  on  segments  2  and  3 
extending  ventrally;  small  lateral  lobes  on  segment  4.  Nephridial 
papillae  on  segments  3,  6-8;  those  of  segment  3  prominent  (anterior 
to  first  setiger).  Thoracic  setigers  11  (12  on  subsp.  pacifica).  Uncini 
begin  on  segment  5  (setiger  2),  single  on  first  six,  in  double  rows  on 
next  eight,  in  single  rows  on  prominent  abdominal  pinnules.  Noto- 
setae  weakly  limbate  (fig.  37,  c).  Pygidium  crenulate.  Females 
with  large  yolky  eggs  in  body  (Point  Barrow,  Sept.  9  and  15,  1948; 
Aug.  17,  Oct.  11,  1949;  Aug.  1,  1950).  Color:  In  life:  Orange,  with 
white  shields,  tentacles  tan.  In  alcohol:  Colorless.  Tube:  With 
thin,  transparent  membranous  lining,  with  very  loose  mud,  bits  of 
sand,  rock,  foraminiferans  (tube  of  specimen  from  Labrador  with 
great  deal  of  loosely  assembled  shell  debris) . 

Remarks. — The  type  of  Leaena  nuda  Moore  from  southwestern 
Alaska  was  examined  and  found  in  poor  condition.  It  has  been  referred 
previously  to  Lanassa  venusta  by  Annenkova  (1937). 

New  records. — Arctic  Alaska:  Point  Barrow  base,  washed  ashore; 
off  Point  Barrow  base,  up  to  12.1  mUes  from  shore,  18.3-75.5  fms.,  on 
bottoms  of  mud,  stones,  and  various  combinations  of  mud,  sand, 
gravel,  pebbles,  stones,  rocks,  large  perforated  rocks,  with  bryozoans, 
shells   (17   stations,   44   specimens).     Canadian  Arctic:    Cove  in 


MARINE    POLYCHAETE    WORMS — ^PETTIBONE  327 

BJQeeland  Bay,  Frobisher  Bay,  Baffin  Island,  14  fms.,  Bartlett,  1942. 
Distribution. — Scattered  records  in  the  Arctic:  Siberian,  Alaskan, 
and  Canadian  Arctic.  Also  Swedish  west  coast,  Danish  waters; 
Labrador;  southwestern  Alaska;  north  Japan  Sea.  In  7-140.5 
fathoms. 

G^nus  Thelepus  Leuckart,  1849 

Thelepua  cincinnatus  (Fabricius,  1780) 

Figure  37,  d 

Amphitrite  cindnnata  Fabricius,  1780,  p.  286. 

Thelepus  circinnaia  Malmgren,  1865,  p.  387,  pi.  22,  fig.  58. 

Thelepus  cincinnatus  Webster  and  Benedict,  1887,  p.  749. — Hessle,  1917,  p.  212. — 
Chamberlin,  1920,  p.  23.— Eliason,  1920,  p.  76.— Fauvel,  1927,  p.  271,  fig. 
95,  i-ra;  1932,  p.  233,  fig.  40;  1934a,  p.  69.— Augener,  1928,  p.  790.— Monro, 
1930,  p.  192;  1936,  p.  182.— Treadwell,  1937,  p.  33.— Friedrich,  1939,  p.  127.— 
Hartman,  1944a,  pp.  336,  343;  1952,  p.  236.— Gorbunov,  1946,  p.  39.— 
Zatsepin,  1948,  p.  154,  pi.  38,  fig.  7.— Wesenberg-Lund,  1950a,  p.  54; 
1950b,  p.  122;  1951,  p.  112. 

Thelepus  hamatus  Moore,  1905b,  p.  856,  pi.  44,  figs.  16-18. — Hartman,  1948, 
p.  44. — Berkeley  and  Berkeley,  1952,  p.  82,  fig.  167. 

Description. — Up  to  200  mm.  long,  10  mm.  wide,  segments  nu- 
merous. Dorsal  surface  rugose,  with  small  glandular  warts  irregularly 
distributed  and  especially  abundant  anteriorly.  Ventral  shields  in- 
distinct, wide,  somewhat  wrinkled.  Buccal  segment  and  prostomium 
forming  a  thick  lower  lip,  a  prominent  horseshoe-shaped  upper  Hp, 
a  semicircular  cephalic  ridge  with  numerous  small  eye-spots;  tentacular 
area  between  upper  lip  and  cephalic  ridge  with  numerous,  rather 
thick  tentacles.  Branchiae  2  pairs  on  segments  2  and  3,  each  formed 
of  numerous,  simple  filaments  arranged  in  parallel  transverse  rows. 
Nephridial  papiUae  on  segments  4-7.  Notosetae  begin  on  segment  3, 
continuing  on  about  40  segments  (or  almost  to  posterior  end).  Un- 
cini  begin  on  segment  5  (setiger  3),  in  single  rows  throughout,  the 
uncinigerous  tori  gradually  transformed  into  projecting  rectangular 
pinnules.  Pygidium  with  crenulate  anal  opening.  Color:  In  life: 
Orange  with  red  branchiae,  tan  tentacles,  with  whitish  glandular 
areas  ventrally  and  laterally.  In  alcohol:  Colorless.  Tube:  Cylin- 
drical, twisted,  with  tough,  transparent,  membranous  lining,  covered 
with  small  pebbles  of  varying  sizes,  with  fragments  of  shells,  bryo- 
zoans,  foraminiferans,  old  worm  tubes,  algae. 

Remarks. — The  tj^pe  of  T.  hamatus  Moore  from  Alaska  was  ex- 
amined and  is  herein  referred  to  T.  cincinnatus;  it  is  a  small  specimen, 
consisting  of  anterior  end  only. 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  up  to  15 
miles  from  shore,  20-123.5  fms,,  on  bottoms  of  mass  of  worm  tubes, 
rocks,  stones,  gravel,  attached  to  stems  of  Tuhularia  (6  stations,  7 
specimens).     West  Greenland:  Godhavn,  off  Hare  Island,  70°20' 


328  PROCEEDINGS    OF   THE   NATIONAL   MUSEUM  vol.108 

N.,  56°  W.,  90  fms.,  Greely  Relief  Expedition,  1884.  One  mile 
northwest  Conical  Rock,  25-60  fms.,  and  north  shore  Wolstenholm, 
13-25  fms.,  Bartlett,  1940.  East  Greenland:  Off  Cape  Hold  with 
Hope,  23-40  fms.,  Bartlett,  1939.  East  Coast  North  America: 
Off  Labrador,  12-30  fms..  Blue  Dolphin  Expedition,  1949;  off  New 
Brunswick,  Nova  Scotia,  Maine,  New  Hampshire,  Massachusetts,  16- 
640  fms.,  U.  S.  Fish  Commission.  West  Coast  North  America: 
Washington  and  Puget  Sounds,  low  water  to  46  fms.,  mud,  shell,  rock, 
Pettibone. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian,  Alaskan, 
and  Canadian  Arctic,  Greenland,  Jan  Mayen,  Spitsbergen,  Franz 
Josef  Land,  Barents  Sea,  Novaya  Zemlya.  Also  Iceland,  Faroes, 
Shetlands,  Norway  to  Portugal,  Azores,  Madeira,  Mediterranean, 
Adriatic;  Labrador  to  Massachusetts;  Bering  Sea  to  Washington; 
Japan;  Indian  Ocean  (Andamans).  Southern  latitudes:  South  Geor- 
gia, South  Orkneys,  South  Shetlands,  Graham  Coast,  Palmer 
Peninsula.     In  low  water  to  1,391  fathoms. 

Genus  Polycirrus  Grube,  1851 
Polycirrus  medusa  Grube,  1855 

Figure  37,  e,  f 

Polycirrus  medusa  Grube,  1855,  p.  120. — Hessle,  1917,  p.  220. — Fauvel,  1927,  p. 
279,  fig.  97,  a-d.— Augener,  1928,  p.  795.— Annenkova,  1934,  p.  322;  1937, 
p.  194;  1938,  p.  210.— Gorbunov,  1946,  p.  39.— Zatsepin,  1948,  p.  154,  pi.  38, 
fig.  5.— Wesenberg-Lund,  1950a,  p.  54;  1950b,  p.  123;  1951,  p.  114. 

Ereutho  smitti  Malmgren,  1865,  p.  391,  pi.  23,  fig.  63. — Webster  and  Benedict, 
1887,  p.  749. 

Polycirrus  sp.,  Hartman,  1948,  p.  45. 

Description. — Length  up  to  70  mm.,  width  6  mm.  Integument 
tesselated.  Large  unpaired  ventral  shield  on  first  few  segments, 
followed  by  about  six  pairs  of  ventral  shields  separated  in  midline. 
Prostomium  and  buccal  segment  forming  lower  lip,  wide  semicircular 
dorsal  cephalic  ridge  (without  eye-spots) ,  large  trilobed  or  undulating 
tentacular  membrane  with  very  numerous  tentacles  on  its  outer  edge, 
the  inner  part  forming  folded  upper  lip;  some  tentacles  long,  slender, 
filiform,  others  thicker,  distinctly  grooved.  Thoracic  setigers  usually 
13  (10-13),  beginning  on  segment  3  (or  2).  Without  thoracic  uncini; 
uncini  begin  on  segment  16  (or  15  or  just  posterior  to  last  setiger), 
uniserial,  few  in  number,  very  small  and  inconspicuous,  on  slightly 
raised  abdominal  pinnules.  Six  pairs  nephridial  papillae  on  segments 
3-8.  Females  with  large,  yolky  eggs  (Point  Barrow,  September  15, 
1948,  and  October  14,  1949).  Color:  In  life:  Red.  In  alcohol: 
Colorless.     Tube  :  Simple  galleries  in  mud  or  sand. 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  up  to  15 
miles  from  shore,  13.3-78.2  fms.,  on  bottoms  of  mud  and  various  com- 


MARINE    POLYCHAETE    WORMS — PETTIBONE  329 

binations  of  mud,  pebbles,  gravel,  stones,  rocks,  large  perforated 
rocks,  with  bryozoans,  worm  tubes  (13  stations,  35  specimens). 
Spitsbergen:  Spitsbergen  Sea,  U.  S.  S.  Alliance,  1881.  Northwest 
Greenland:  One  mile  northwest  Conical  Rock,  25-60  fms.,  Bartlett, 
1940.  Bering  Sea:  St.  Paul  Island,  Pribilofs,  Palmer,  1890.  East 
Coast  North  America:  Off  Labrador,  45  fms.,  silt,  Blue  Dolphin 
Expedition,  1949.  West  Coast  North  America:  Pavlof  Bay, 
Alaska,  150  fms.,  Alaska  King  Crab  Investigation,  1940;  Washington 
Sound,  12-46  fms.,  mud  and  mussels,  Pettibone. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian  and 
Alaskan  Arctic,  Greenland,  Spitsbergen,  White  Sea,  Novaya  Zemlya, 
Kara  Sea.  Also  Iceland,  Faroes,  Swedish  west  coast  to  France, 
Mediterranean;  Labrador  to  Maine;  Bering  Sea  to  Washington; 
Okhotsk  Sea  to  north  Japan  Sea.     In  low  water  to  889  fathoms. 

Genus  Trichohranchus  Malmgren,  1865 
Trtchobranchus  glacialis  Malmgren,  1865 

Figure  37,  g-i 

Trichohranchus  glacialis  Malmgren,  1865,  p.  395,  pi.  24,  fig.  65. — Webster  and 
Benedict,  1887,  p.  750.— Ehlers,  1913,  p.  566.— Hessle,  1917,  p.  131.— 
Fauvel,  1927,  p.  288,  fig.lOO,a-h.— Augener,  1928,  p.  792.— Annenkova, 
1937,  p.  190;  1938,  p.  202.— Zatsepin,  1948,  p.  153,  pi.  38,  fig.  1.— Wesenberg- 
Lund,  1950a,  p.  55;  1950b,  p.  125;  1951,  p.  115.— Berkeley  and  Berkeley, 
1952,  p.  76,  figs.  154,  155. 

Trichohranchus  glacialis  var.  antarcticus  Hessle,  1917,  p.  132. — Hartman,  1952, 
p.  233. 

Description. — Length  up  to  30  mm.,  width  3  mm.,  segments  60-70. 
Buccal  segment  and  prostomium  form  thick,  tesselated,  inflated  lower 
lip  connected  laterally  with  pair  of  projecting,  rounded,  flattened  lobes 
(ventral  to  mass  of  tentacles),  cephalic  ridge  with  numerous  black 
eyespots  (absent  in  var.  antarcticus) ,  undulating  tentacular  membrane 
with  very  numerous  tentacles,  and  folded  upper  lip;  some  tentacles 
larger,  distinctly  grooved;  others  smaller,  filiform.  Three  pairs 
branchiae  on  segments  2-4,  each  composed  of  single  long  filament. 
With  slightly  projecting  lateral  lobes  on  branchial  segments.  Thoracic 
setigers  and  uncinigers  15,  beginning  on  segment  6;  notosetae  limbate, 
with  smooth  capillary  tips;  neuropodial  uncini  aciculiform  (crotchets). 
Abdominal  segments  with  avicular  uncini  in  single  rows  on  triangular 
projecting  pinnules.  Pygidium  with  anal  opening  crenulate.  Color- 
less in  alcohol.     Tube  membranous,  encrusted  with  mud  or  fine  sand. 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  up  to  5 
miles  from  shore,  27-49  fms.,  on  various  combinations  of  mud,  gravel, 
stones,  rocks,  shells  (5  stations,  15  specimens).  East  Coast  North 
America:  Cape  Cod  Bay,  Massachusetts,  27-118  fms.,  U.  S.  Fish 
Commission. 


330  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  los 

Distribution. — Widely'^distributed  in  the  Arctic:  Alaskan  Arctic, 
Greenland,  Spitsbergen,  Novaya  Zemlya,  Kara  Sea.  Also  Iceland, 
Faroes,  Swedish  west  coast  to  Spain,  Canary  Islands,  Mediterranean; 
Maine  to  Massachusetts;  British  Columbia;  north  Japan  Sea.  Var. 
antarcticus:  Magellan  Straits,  South  Georgia,  Ross  Island,  Kaiser 
Wilhehn  II  Land.     In  3-1,517  fathoms. 

Genus  Terehellides  M.  Sars,  1835 
Terebellides  stroetnii  M.  Sars,  1835 

FiQUEB  37,  j-m 

Terebellides  stroemii  M.  Sars,  1835,  p.  48,  pi.  13,  fig.  31. — Malmgren,  1865,  p 
396,  pi.  20,  fig.  48.— Webster  and  Benedict,  1887,  p.  750.— Moore,  1903,  p 
478;  1908,  p.  352;  1923,  p.  199.— Hessle,  1917,  p.  137.— Chamberlin,  1920 
p.  23.— Eliason,  1920,  p.  72.— Fauvel,  1927,  p.  291,  fig.  100,  i-q;  1932,  p 
234;  1947,  p.  79,  fig.  76,  f-i.— Augener,  1928,  p.  797.— Gustafson,  1936,  p 
9.— Annenkova,  1937,  p.  190;  1938,  p.  202.— Treadwell,  1937,  p.  35.— Okuda, 
1938b,  p.  102.— Friedrich,  1939,  p.  127.— Berkeley  and  Berkeley,  1943,  p 
130;  1944,  p.  5;  1952,  p.  75,  figs.  152,  153.— Hartman,  1944a,  pp.  336,  343 
1944d,  p.  24;  1951,  p.  113.— Thorson,  1946,  p.  124,  fig.  69.— Gorbunov,  1946 
p.  39.— Zatsepin,  1948,  p.  153,  pi.  38,  fig.  3.— St0p-Bowitz,  1948c,  p.  68.— 
Wesenberg-Lund,  1949,  p.  355;  1950a,  p.  55;  1950b,  p.  126;  1951,  p.  115,  fig 
11. — Hartman  and  Reish,  1950,  p.  44. 

Terebellides  stroemii  var.  japonica  Moore,  1903,  p.  478;  1908,  p.  352;  1923,  p.  200 

Description. — Length  up  to  75  mm.,  width  8  mm.,  segments  50-60 
Prostomium  and  buccal  segment  forming  large,  transverse,  crescent- 
shaped  plate  below  mouth,  large  and  undulating  tentacular  membrane 
with  numerous,  short,  grooved  tentacles,  folded  upper  lip;  dorsaUy 
without  cephalic  ridge  or  eye-spots.  Segment  2  short,  achaetous. 
Single  branchia  formed  of  large  cylindrical  trunk  dorsally  on  segment 
3  (first  setiger)  and  four  pectinate  lobes,  frequently  interlocked  giving 
appearance  of  more  or  less  solid  mass.  With  somewhat  developed 
lateral  lobes  extending  ventraUy  on  segments  2-7  (larger  on  segments 
3-6).  Thoracic  setigers  18,  begin  on  segment  3.  Uncini  begin  on 
segment  8  (setiger  6),  those  on  first  unciniger  consisting  of  crotchets 
with  long  manubrium  and  bent,  pointed  tips,  those  on  rest  of  thoracic 
segments  long,  aciculiform,  with  large  fang  and  few  denticles;  may  be 
crowded  in  more  than  one  row  in  large  specimens.  Abdominal 
uncini  short,  avicular,  pectiniform,  in  single  rows  on  projecting  tri- 
angular pinnules.  Prominent  paired  nephridial  papillae  on  segment  3, 
smaller  ones  on  segments  6  and  7.  Pygidium  with  terminal  anus,  with 
sides  crenulate.  Female  with  large  yolky  eggs  (ripe  eggs  and  sperm, 
September  26,  1949).  Color:  In  life:  Body  orange,  branchia  orange- 
red,  tentacles  tan.  In  alcohol:  Colorless,  with  smooth  iridescent 
surface. 

Parasites. — One  of  the  87  specimens  had  the  parasitic  copepod 
Saccopsis  terebellidis  Levinsen  (identified  by  P.  L.  lUg)  attached  to 


MARINE    POLYCHAETE    WORMS — PETTIBONE 


331 


Figure  37. — Terebellidae :  a,  Leaena  abranchiata,  lateral  view  anterior  end;  b,  same,  notoseta; 
c,  Lanassa  venusta,  notoseta;  d,  Thelepus  cincinnatus,  avicular  uncinus;  e,  Polycirrus 
medusa,  dorsal  view  anterior  end;/,  same,  ventral  view  anterior  end;  g,  Trichobranchus 
glacialis,  dorsal  view  anterior  end;  h,  same,  thoracic  aciculiform  uncinus;  i,  same,  abdom- 
inal avicular  uncinus;  ;',  Terebellides  stroemii,  lateral  view  anterior  end;  k,  same,  crotchet 
from  first  unciniger;  /,  same,  thoracic  aciculiform  uncinus;  m,  same,  abdominal  avicular 
uncinus.     (For  explanation  of  symbols,  see  p.  210.) 


261112—54- 


332  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  103 

anterior  part  in  the  region  of  the  branchia.  The  same  polychaete  and 
copepod  species  were  found  in  Iceland  waters  and  figured  by  Wesen- 
berg-Lund  (1951,  p.  117,  fig.  11). 

New  records. — Arctic  Alaska:  West  side  Elson  Lagoon  near  Point 
Barrow,  1.2  fms,;  Eluitkak  Pass,  Elson  Lagoon;  Point  Barrow  base, 
washed  ashore;  off  Point  Barrow  base,  up  to  15  miles  from  shore,  18.3- 
123.5  fms.,  on  bottoms  of  mud,  stones,  mass  of  worm  tubes,  and  various 
combinations  of  mud,  sand,  pebbles,  gravel,  stones,  rocks,  large  per- 
forated rocks,  with  shells,  bryozoans,  worm  tubes  (26  stations,  87 
specimens).  West  Coast  North  America:  Washington  and  Puget 
Sounds,  10-165  fms.,  Pettibone.  East  Coast  North  America:  Off 
Labrador,  5-65  fms..  Blue  Dolphin  Expeditions,  1949,  1950;  off  Nova 
Scotia,  Maine,  Massachusetts,  Rhode  Island,  Long  Island  Sound, 
6-368  fms.,  U.  S.  Fish  Commission. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian,  Alaskan, 
and  Canadian  Arctic,  Greenland,  Jan  Mayen,  Spitsbergen,  Franz 
Josef  Land,  Barents  Sea,  White  Sea,  Novaya  Zemlya,  Kara  Sea.  Also 
Iceland,  Faroes,  Norway  to  Portugal,  Mediterranean,  Adriatic,  Black 
Sea;  Hudson  Bay  to  Long  Island  Sound,  Gulf  of  Mexico,  West  Indies; 
Bering  Sea  to  southern  California,  Panamd,  Venezuela;  north  Japan 
Sea  to  Japan;  Iranian  Gulf;  Indian  Ocean;  South  Africa,  Bouvet 
Island,  Kerguelen,  New  Caledonia.    In  low  water  to  1,611  fathoms. 

Family  Sabellidae 

Body  cylindrical,  tapered  posteriorly.  Prostomium  indistinct. 
Buccal  region  with  mouth  terminal,  with  dorsal  lip,  two  ventral  lips 
or  vesicular  bulbs,  two  fleshy,  membranous  or  filiform  palps  of  variable 
length.  Branchiae  form  conspicuous,  often  highly  colored  terminal 
funnellike  plume  surrounding  mouth;  plume  formed  of  two  semi- 
circular or  spiral  lobes  bearing  numerous  filaments  each  with  two  rows 
of  ciliated  barbules  (fig.  38,  a).  Branchiae  without  operculum.  A 
collarette  more  or  less  developed  on  first  segment.  Body  divided 
into  two  regions:  (1)  thoracic;  few  segments  (4-12),  with  dorsal  bun- 
dles of  capillary  setae  and  ventral  uncinigerous  tori  (tori  lacking  in 
Myxicola);  without  thoracic  membrane;  (2)  abdominal;  with  inversion 


Figure  38. — Sabellidae:  a,  Sabella  crassicornis,  lateral  view  anterior  end;  b,  same,  tip 
of  branchial  filament;  c,  same,  thoracic  notosetae;  d,  same,  dorsal  view  pygidium;^,  same, 
thoracic  hoe-like  seta  from  ventral  torus;/,  same,  thoracic  avicular  uncinus  from  ventral 
torus;  g,  same,  ventral  view  collarette;  h,  same,  lateral  view;  i,  same,  dorsal  view;  /,  Pot- 
amilla  neglecta,  dorsal  view  collarette;  k,  same,  lateral  view;  /,  same,  ventral  view;  m, 
same,  dorsal  thoracic  limbate  seta;  n,  same,  dorsal  thoracic  spatulate  setae;  o,  Potamilla 
reniformis,  ventral  view  collarette;  p,  same,  lateral  view;  q,  same,  dorsal  view;  r,  same, 
portion  of  branchial  filament;  s,  same,  thoracic  avicular  uncinus  from  ventral  torus;  t, 
same,  thoracic  hoe-like  seta  from  ventral  torus;  «,  same,  tip  of  tube,  rolled  at  free  end. 
(For  explanation  of  symbols,  see  p.  210.) 


MARINE    POLYCHAETE    WORMS — PETTIBONE 

I 


333 


Figure  38. — For  explanation  see  facing  page. 


334  PROCEEDINGS    OF   THE    NATIONAL    MUSEUM  vol.  los 

of  setae,  dorsal  uncinigerous  tori  and  ventral  capillary  setae.  A 
median  ciliated  faecal  groove  runs  length  of  body,  ventral  in  abdom- 
inal region  and  crossing  to  dorsal  surface  in  thoracic  region.  Tube 
cylindrical,  permanent  or  more  or  less  transitory,  gelatinous,  mem- 
branous, or  horny,  covered  or  not  with  mud,  sand,  gravel,  debris  of 
shells. 

Represented  by  five  genera  and  seven  species  at  Point  Barrow. 
All  the  genera  have  large  branchial  plumes  composed  of  two  similar 
semicircular  lobes;  branchial  filaments  without  large  subterminal 
eyes,  without  dorsal  appendages.  First  setigerous  segments  with 
limbate  notosetae  only;  neurosetae  begin  on  setiger  2.  Abdominal 
region  with  numerous  segments. 

Key  to  the  genera  of  SaheUidae  from  Point  Barrow 

1.  Ventral  tori  of  thorax  with  row  of  uncini  of  the  avicular  form  (fig.  38,  /;  in 

Sabella  and  Potamilla  with  additional  row  of  hoelike  setae,  fig.  38,  t) .     Col- 
larette well  developed.     Branchial  filaments  with  or  without  eyes,  united 

at  base  only,  with  palmar  membrane  poorly  developed  or  lacking 2 

Ventral  tori  of  thorax  with  crotchets  with  long  manubrium  (fig.  39,  t;  without 
tori  in  Myxicola).  Branchial  filaments  without  eyes,  united  along  a  large 
part  of  their  length  by  a  well-developed  palmar  membrane  (fig.  39,  w)  —  3 

2.  Dorsal  thoracic  setae  all  limbate,  some  long  with  straight  borders,  others  short 

with  wide  borders  (fig.  38,  c) Sabella   (p.  334) 

Dorsal  thoracic  setae  of  two  kinds,  some  limbate,  others  spatulate  (fig.  38,  m,  n) 

Potamilla  (p.  335) 

3.  Uncinigerous  tori  projecting,  with  short,  lateral  rows  of  uncini.     Collarette 

well  developed  (fig.  39,  a-c).     Dorsal  thoracic  setae  of  two  kinds,  limbate 

and  spatulate  (fig.  39,  /,  g) 4 

Without  projecting  uncinigerous  tori,  abdominal  uncini  in  almost  complete 
transverse  band  (fig.  39,  p).  Collarette  poorly  developed,  represented  by 
a  triangular  ventral  lobe  (fig.  39,  o).  Dorsal  thoracic  setae  all  capillary, 
limbate Myxicola  (p.  340) 

4.  Posterior  segments  without  ventral  depression Chone  (p.  337) 

Posterior  segments  about  10,  with  ventral  suckerlike  disc,  large  ventral  de- 
pression with  flared  sides  (fig.  39,  n) Euchoiie  (p.  339) 

Genus  Sabella  Linn^,  1767  (sensu  Malmgren,  1865) 

Sabella  crassicornis  Sars,  18.51 

Figure  38,  a-i 

Sabella  crassicornis  Sars,  1851,  p.  202.— Malmgren,  1865,  p.  399,  pi.  27,  fig.  83.— 

Moore,  1909b,  p.  144.— Johansson,  1927,  p.  119.— Hartman,  1942a,  p.  78; 

1948,  p.  46.— Berkeley  and  Berkeley,  1943,  p.  130;  1952,  p.  114,  figs.  236, 

237.— Zatsepiu,  1948,  p.  161,  pi.  39,  fig.  4. 
Sabella  fabricii  Kroyer,  1856,  p.  20.— Fauvel,  1927,  p.  300,  fig.  103,  a-g.— Augener, 

1928,  p.  800.— Annenkova,  1934,  p.  322;  1937,  p.  195;  1938,  p.  211.— Wesen- 

berg-Lund,  1950b,  p.  128;  1951,  p.  118. 
Sabella  spetsbergensis  Malmgren,  1865,  p.  399,  pi.  29,  fig.  93. 
Sabella  spitzbergensis  Webster  and  Benedict,  1887,  p.  750. 


MARINE    POLYCHAETE    WORMS — PETTIBONE  335 

Description. — Length  20-80  mm.,  width  3-4  mm.  Collarette 
widely  separated  middorsally,  with  midventral  slit  and  lateral  notches, 
resulting  in  4-lobed  structm-e;  ventral  lobes  closely  approximated, 
may  be  deflected.  Two  branchial  lobes  each  with  about  16  filaments 
(15-35),  with  short  tapering  tips.  Branchial  filaments  with  usually 
four  to  six  (2-8)  pairs  of  eyes  (located  in  color  bands  of  filaments  and 
may  easily  be  overlooked).  Thoracic  setigers  usually  8  (7-9).  Abdo- 
men with  one  or  two  pairs  of  eye-spots  between  parapodial  rami,  well 
developed  toward  posterior  end.  Pj^gidium  with  pair  of  bulbous 
lobes  with  groups  of  eye-spots  on  dorsolateral  sides.  Color:  In 
life :  Body  flesh  color,  with  branchial  filaments  banded  rusty  red,  4-7 
transverse  bands  per  filament.  In  alcohol :  Body  colorless  with  red- 
dish purple  bands  on  branchial  filaments.  Tube:  Cylindrical,  free 
end  flexible,  covered  with  thin,  smooth  layer  of  mud;  embedded  part 
transparent,  horny,  rigid,  covered  with  sand  grains  and  foreign 
material. 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  up  to  15 
miles  from  shore,  36-123.5  fms.,  on  bottoms  of  stones,  mass  of  worm 
tubes,  and  various  combinations  of  gravel,  stones,  rocks,  large  per- 
forated rocks,  with  worm  tubes  (8  stations,  15  specimens).  Kam- 
chatka: Petropavlovsk,  Grevnitzky,  1888.  Alaska:  New  Harbor, 
Unga  Island,  under  stones,  Dall,  1872;  Albatross  Sta.  2847,  55°01'  N., 
160°18'  W.,  48  fms.,  and  station  at  Kodiak,  1888.  Canadian  Arctic: 
Foxe  Basin,  25-31  fms.,  Bartlett,  1927.  East  Coast  North  America: 
Off  Labrador,  5-6  fms..  Blue  Dolphin  Expedition,  1949;  Bay  of  Fundy, 
Maine,  Massachusetts,  15  fms.,  U.  S.  Fish  Commission.  Central 
America:  Golfo  Dulce,  west  Costa  Rica,  M.  Valerio. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian,  Alaskan, 
and  Canadian  Arctic,  Greenland,  Spitsbergen,  Novaya  Zemlya,  Kara 
Sea.  Also  Iceland,  Faroes,  Norway  to  France,  Mediterranean; 
Hudson  Bay  to  Massachusetts;  Bering  Sea  to  California,  Central 
America  (Costa  Rica);  north  Japan  Sea  to  Japan.  In  low  water  to 
230  fathoms. 

Genus  Potaniilla  Malmgren,  1865 

Key  to  the  species  of  Potamilla  from  Point  Barrow 

1.  Without  branchial  eyes P.  neglecta 

With  compound  eyes  in  single  rows  on  branchial  filaments,  0-8  per  filament 
(fig.  38,  r) P.  reniformis 

Potamilla  neglecta  (Sars,  1851) 

Figure  38,  j-n 

Sabella  neglecta  Sars,  1851,  p.  203. 

Potamilla  neglecta  Malmgren,  1865,  p.  401,  pi.  27,  fig.  84. — Webster  and  Benedict, 
1884,  p.  736.— Moore,  1909b,  p.  145;  1923,  p.  242.— Johansson,  1927,  p.  143.— 


336  PROCEEDINGS    OF   THE    NATIONAL   MUSEUM  vol.  103 

Augener,    1928,    p.    801.— Hartman,    1942a,    p.    81;   1944a,    pp.   336,  343; 

1948,  p.  46.— Zatsepin,  1948,  p.  162,  pi.  39,  fig.  7.— Wesenberg-Lund,  1950a, 

p.  56;  1950b,  p.  128;  1951,  p.  119.— Berkeley  and  Berkeley,  1952,  p.  116,  fig. 

238. 
Potamilla  torelli  Malmgren,  1865,  p.  402;  1867,  p.  114,  pi.  13,  fig.  76.— Ehlers, 

1913,  p.  575.— Fauvel,  1927,  p.  310,  fig.  107,  m-s;  1934a,  p.  71.— Annenkova, 

1934,  p.  322;  1937,  p.  195;  1938,  p.  213.— Wesenberg-Lund,  1951,  p.  121. 
Potamilla  acuminata  Moore  and  Bush,  1904,  p.  159,  pi.  11,  figs.  3-6;  pi.  12,  fig. 

41. 

Description. — Length  30-84  mm.,  1-3  mm.  wide.  Collarette  widely- 
separated  dorsally,  sloping  ventrally,  with  midventral  slit,  entire 
laterally,  resulting  in  a  2-lobed  structure;  ventral  lobes  triangular, 
may  be  deflected.  Branchial  lobes  each  with  about  16  filaments 
(6-20),  with  short,  naked  distal  tips,  without  eyes.  Thoracic  setigers 
usually  8  (5-8).  Pygidium  with  pair  of  bulbous  lobes  with  dark  eye- 
spots.  Color:  In  alcohol:  Body  colorless,  branchiae  with  two  or 
three  difiFused  reddish  brown  bands.  Tube:  Horny,  transparent, 
more  or  less  covered  with  sand  or  mud;  smaller  tubes  with  rather 
uniform  layer  of  small  sand  grains;  larger  tubes  may  be  encrusted 
with  living  barnacles,  bryozoans,  hydroids,  foraminiferans,  amphipod 
tubes.  Large  yolky  eggs  in  a  single  layer,  pressed  close  to  wall, 
about  one-third  way  down  in  tube;  a  transparent,  thin  membrane 
laid  down  between  eggs  and  worm  (large  developing  yellow  eggs 
found  in  tube,  September  6,  1949,  dredged  in  36  fathoms.  Point 
Barrow) . 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  up  to  12.1 
miles  from  shore,  21.7-123.5  fms.,  on  bottoms  of  mass  of  worm  tubes 
and  pebbles,  gravel,  rocks,  large  stones  (7  stations,  24  specimens). 
Bering  Sea:  Albatross  Sta.  3496,  56°32'  N.,  169°45'  W.,  41  fms., 
1893;  St.  Paul  Island,  Pribilofs,  Hahn,  1911.  Alaska:  Albatross 
Sta.,  Kodiak,  1888.  West  Greenland:  Off  Hare  Island,  70°20'  N., 
56°  W.,  90  fms.,  U.  S.  S.  Alert,  1884.  East  Coast  North  America: 
Off  Labrador,  125  fms.,  Blue  Dolphin  Expedition,  1949;  off  Maine, 
Massachusetts,  31-130  fms.,  U.  S.  Fish  Commission. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian  and 
Alaskan  Arctic,  Greenland,  Spitsbergen.  Also  Iceland,  Faroes,  Nor- 
way to  France,  Madeira,  Mediterranean,  Adriatic,  Cape  Verde 
Islands;  Labrador  to  Massachusetts;  Bering  Sea  to  California;  north 
Japan  Sea  to  Japan;  Antarctic.     In  low  water  to  1,044  fathoms. 

Potamilla  reniformis  (Leuckart,  1849) 

Figure  38,  o-u 

Sabella  reniformis  Leuckart,  1849,  p.  183,  pi.  3,  fig.  8. 

Potamilla  reniformis  Malmgren,  1867,  p.  114,  pi.  13,  fig.  77. — Webster  and  Bene- 
dict, 1884,  p.  736;  1887,  p.  750.— Johansson,  1927,  p.   142.— Fauvel,   1927, 


MARINE    POLYCHAETE    WORMS — PETTIBONE  337 

p.  309,  fig.  107,  a-1.— Okuda,  1937b,  p.  61.— Annenkova,  1934,  p.  322;   1937, 

p.  195;  1938,  p.  212.— Zatsepin,  1948,  p.  161,  pi.  39,  fig.  6.— Wesenberg-Lund, 

1950a,  p.  57;  1950b,  p.  129;  1951,  p.  120. 
Pseudopoiamilla  intermedia  Moore,  1905c,  p.  562,  pi.  37,  figs.  15-22;  1908,  p.  359. — 

Hartman,   1938c,  p.  25,  pi.  2,  fig.  8;  pi.  3,  figs.  1-4;  1948,  p.  46.— Rioja, 

1941,  p.  732. 
Pseudopotamilla  reniformis  Hartman,  1944a,  pp.  336,  343,  pi.  21,  figs.  3,  6;  1945, 

p.  47.— Berkeley  and  Berkeley,  1952,  p.  116,  fig.  239. 

Description. — Length  80-100  mm.,  width  1-2  mm.  Collarette 
with  middorsal  depression,  deeply  notched  dorsolaterally  (at  level  of 
notopodia),  with  midventral  slit,  resulting  in  a  4-lobed  structm-e  of 
pair  of  rounded  or  triangular  dorsal  lobes  and  pair  of  rounded  ventral 
lobes  which  may  be  deflected.  Branchial  lobes  each  with  about  10 
filaments  (5-15).  At  least  some  of  branchial  filaments  with  large 
compound  eyes,  usually  0-3  per  filament  (0-8),  eyes  may  be  variable 
in  size  or  subequal,  deep  reddish  in  alcohol.  Thoracic  setigers  usually 
10  (7-14).  Pygidium  with  pair  of  bulbous  lobes  with  eye-spots. 
Color:  In  alcohol:  Body  colorless,  branchiae  with  diffused  reddish 
brown  bands  in  region  of  eyes.  Tube:  Somewhat  twisted,  horny, 
translucent  or  opaque,  more  or  less  covered  with  sand  or  mud,  the 
free  end  usually  flattened,  rolled  scroll-like  when  animal  pulls  inside. 

New  records. — Arctic  Alaska:  Off  Point  Barrow  base,  up  to  7.4 
miles  from  shore,  21.7-54.6  fms.,  on  bottoms  of  pebbles,  gravel, 
rocks,  in  holes  of  large  stones  (2  stations,  15  specimens).  Bering 
Sea:  Anchorage,  Cape  Etoliu,  Nunivak  Island,  8  fms.,  Dall,  1874. 
Southeastern  Alaska:  Lituya  Bay,  6-9  fms.,  Dall,  1874.  East 
Coast  North  America:  Off  Nova  Scotia,  Maine,  New  Hampshire, 
Massachusetts,  Rhode  Island,  Delaware,  10-317  fms.,  U.  S.  Fish 
Commission. 

Distribution. — Scattered  records  in  the  Arctic:  Arctic  Alaska, 
Greenland.  Also  Iceland,  Swedish  west  coast  to  France,  Mediter- 
ranean; Nova  Scotia  to  North  Carolina;  Bering  Sea  to  British  Co- 
lumbia, Mexico;  north  Japan  Sea  to  Japan.  In  low  water  to  317 
fathoms. 

Genus  Chone  Kroyer,  1856 

Both  species  have  the  segments  biannulate.  Collarette  deeply 
incised  middorsally,  forming  two  longitudinal-folded  brackets,  with 
sides  smooth;  without  midventral  or  lateral  notches  (fig.  39,  a-c). 
Branchial  filaments  united  on  great  part  of  their  length  by  palmar 
membrane  (fig.  39,  m).  Thoracic  setigers  8.  Thoracic  dorsal  setae 
of  three  kinds — limbate  capillary  setae,  fine  bayonette  setae,  and 
spatulate  setae  with  asymmetrical  mucronate  tips  or  gently  rounded 
(fig.  39,f-h).  Pygidium  with  anus  terminal,  with  rounded  to  conical 
bulbous  dorsal  valve  (fig.  39,  d). 


338  PROCEEDINGS    OF   THE    NATIONAL    MUSEUM  vol.  103 

Key  to  the  species  of  Chone  from  Point  Barrow 

1.  Collarette  nearly  straight  (fig.  39,  b).  Free  end  of  branchial  filaments  flat- 
tened, foliaceous,  widely  limbate  (fig.  39,  e)_- -_C.  infuudibuliformis 

Collarette  oblique,  longer  on  ventral  side  (fig.  39,  k).     Free  end  of  branchial 
filaments  with  long  filiform  tips  (fig.  39,  I) C.  duneri 

Chone  infundibuliformia  Kroyer,  1856 

Figure  39,  a-j 

Chone  infundibuliformis  Kroyer,  1856,  p.  33. — Malmgren,  1865,  p.  404,  pi.  28, 
fig.  87.— Th6el,  1879,  p.  66.— Wir6n,  1883,  p.  422.— Fauvel,  1927,  p.  334, 
fig.  116,  a-o.— Augener,  1928,  p.  806.— Annenkova,  1934,  p.  322;  1937, 
p.  196;  1938,  p.  215.— Hartman,  1942b,  p.  136;  1944a,  pp.  334,  336  (not  pi. 
20,  fig.  5),  pi.  21,  fig.  7.— Berkeley  and  Berkeley,  1943,  p.  130;  1952,  p.  123, 
figs.  252,  253.— Gorbunov,  1946,  p.  39.— Zatsepin,  1948,  p.  164,  pi.  39, 
fig.  13.— Wesenberg-Lund,  1950a,  p.  58;  1950b,  p.  131;  1951,  p.  123. 

Chone  gracilis  Moore,  1906b,  p.  257,  pi.  12,  figs.  62-66. — Berkeley  and  Berkeley, 
1952,  p.  123,  fig.  254. 

Description. — Length  30  mm.,  width  2.5  mm.  (up  to  120  mm.  long, 
6  mm.  wide — Fauvel,  1927).  Collarette  nearly  straight  laterally. 
Branchial  filaments  about  15  (10-36),  with  tips  more  or  less  foliaceous, 
edged  by  transparent  border  (prolongations  of  palmar  membrane). 
Color:  In  life:  Variable — flesh  color,  branchiae  orange  spotted  with 
white;  olive  green  with  distal  half  of  branchiae  brick  red,  basal  half 
red  and  olive,  chalk  white  on  outer  sides  of  bases  of  branchiae.  In 
alcohol:  Colorless,  gra3dsh,  or  tannish.  Tube:  Membranous,  en- 
crusted with  sand,  mud,  or  pebbles  of  variable  sizes,  foraminiferans. 

New  records. — Arctic  Alaska:  Eluitkak  Pass,  Elson  Lagoon  near 
Point  Barrow,  6.6  fms.;  off  Point  Barrow  base,  up  to  7.5  miles  from 
shore,  20^9  fms.,  on  bottoms  of  mud,  stones,  and  various  combina- 
tions of  mud,  sand,  gravel,  stones,  rocks,  large  perforated  rocks,  shells 
(12  stations,  32  specimens).  Canadian  Arctic:  East  side  Cobourg 
Island,  Baffin  Bay,  75°  40'  N.,  78°  40'  W.,  Bartlett,  1935.  North- 
west Greenland:  1  mile  northwest  Conical  Rock,  Bartlett,  1940. 
East  Greenland:  Off  Cape  Hold  with  Hope,  23-40  fms.,  Bartlett, 
1939.  Spitsbergen:  Spitsbergen  Sea,  U.S.S.  ^Z/iance,  1881.  Bering 
Sea:  St.  Paul  Island,  Pribilofs,  Palmer,  1890;  St.  George  Island, 
Pribilofs,  Hanna,  1913;  Albatross  Sta.  3232,  58°  30'  N.,  157°  34'  W., 
10.5  fms.,  1890,  Sta.  3233,  58°  23'  N.,  157°  42'  W.,  7  fms.,  1890,  and 
Sta.  3289,  56°  44'  N.,  159°  16'  W.,  16  fms.,  1890;  Kiska,  Aleutians, 
Dall.  West  Coast  North  America:  Strait  of  Juan  de  Fuca,  Wash- 
ington, 40  fms.,  mud,  Pettibone.  East  Coast  North  America:  Off 
Nova  Scotia,  Maine,  Massachusetts,  Rhode  Island,  10-206  fms.,  U. 
S.  Fish  Commission. 

Distribution. — Widely  distributed  in  the  Arctic:  Siberian,  Alaskan, 
and  Canadian  Arctic,  Greenland,  Spitsbergen,  Novaya  Zemlya,  Kara 


MARINE    POLYCHAETE    WORMS — PETTIBONE  339 

Sea.  Also  Iceland,  Faroes,  Shetlands,  Scandinavian  coast  to  Danish 
waters;  Hudson  Bay  to  Rhode  Island;  Bering  Sea  to  California;  north 
Japan  Sea.     In  low  water  to  1,955  fathoms. 

Chone  dun^ri  Malmgren,  1867 

Figure  39,  k,  I 

Chone  dunSri  Malmgren,  1867,  p.  116,  pi.  13,  fig.  75.— Wir^n,  1883,  p.  422.— 
Fauvel,  1927,  p.  336,  fig.  117,  1-r;  1934a,  p.  74.— Augener,  1928,  p.  807.— 
Monro,  1936,  p.  189.— Treadwell,  1937,  p.  35.— Berkeley  and  Berkeley,  1944, 
p.  5. — Zatsepin,  1948,  p.  164,  pi.  39,  fig.  15. — Wesenberg-Lund,  1950a,  p. 
58;  1950b,  p.  130;  1951,  p.  123.— Hartman,  1951,  p.  117. 

Description. — Length  20-35  mm.,  width  1.5-2.5  mm.  Collarette 
oblique,  longer  on  ventral  side.  Branchial  filaments  about  10  (5-11), 
with  long,  tapered  tips.  Color:  In  life:  Branchiae  yellow.  In  alco- 
hol: Colorless  or  tannish.  Tube:  Thin,  membranous,  encrusted  with 
sand. 

New  records. — ^ Arctic  Alaska:  Off  Point  Barrow  base,  up  to  7.5 
miles  from  shore,  13.3-49  fms.,  on  bottoms  of  mud  and  various  com- 
binations of  mud,  sand,  gravel,  stones,  rocks,  large  perforated  rocks, 
shells  (9  stations,  21  specimens).  Canadian  Arctic:  Baffin  Island, 
66°  43'  N.,  80°  07'  W.,  Bartlett,  1927.  East  Coast  North  America: 
Off  Labrador,  8  fms.,  mud.  Blue  Dolphin  Expedition,  1949. 

Distribution. — Widely  distributed  in  the  Arctic:  Alaskan  and  Ca- 
nadian Arctic,  Greenland,  Spitsbergen,  Novaya  Zemlya,  Kara  Sea. 
Also  Iceland,  Faroes,  Norway  to  North  Sea,  Madeira,  Mediterranean, 
Adriatic;  Labrador;  Florida;  west  coast  South  America  (Perii).  In 
8-889  fathoms. 

Genus  Euchone  Malmgren,  1865 
Euchone  analis  (Kroyer,  1856) 

Figure  39,  m,  n 

Sabella  analis  KrSyer,  1856,  p.  17. 

Euchone  analis  Malmgren,  1865,  p.  406,  pi.  28,  fig.  88.— Th6el,  1879,  p.  65.— 
Wir6n,  1883,  p.  423.— Chamberlin,  1920,  p.  27.— Augener,  1928,  p.  804.— 
Annenkova,  1937,  p.  196;  1938,  p.  216.— Berkeley  and  Berkeley,  1943,  p. 
130;  1952,  p.  121,  figs.  250,  251.— Gorbunov,  1946,  p.  39.— Zatsepin,  1948, 
p.  163,  pi.  39,  fig.  12.— Wesenberg-Lund,  1950a,  p.  59;  1950b,  p.  132;  1951, 
p.  124. 

Description. — Length  19-28  mm.,  width  1.8  mm.  (up  to  60  mm. 
long — Augener,  1928).  Segments  biaimulate  and,  with  the  ventral 
faecal  groove,  forming  four  large  areas  per  segment.  Collarette 
with  opening  dorsally,  nearly  straight  and  entire  laterally,  with 
shallow  midventral  incision.  Branchial  filaments  about  10  per  lobe 
(up  to  14),  united  through  a  great  part  of  their  length